Volume 36 Number 4 April1974
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Marine Fisheries
REVIEW
National Oceanic and Atmospheric Administration • National Marine Fisheries Service
SPECIAL NUMBBt
1HE CAUFORNIA GRAf WHALE
Volume 36 Number 4 April 1974
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Marine Fisheries
REVIEW
National Oceanic and Atmospheric Administration • National Marine Fisheries Service
SPECIAL NUMBER
THE CAUFORNIA GRAY WHALE
Marine Fisheries Review
Vol.36, No. 4
April 1974
CONTENTS
Articles
1 Preface. W. E. Evans
2 Introductory Remarks. Brian J. Rothschild
3 Respiration and Metabolism in Two Baleen Whale Calves, Eric A. Wahren-
brock. Gary F. Maruschak. Robert Eisner, and David W. Kenney
9 Ballistocardiography as a Technique for Comparative Physiology. N. Ty
Smith and Eric A. Wahrenbrock
15 Investigation of Blubber Thickness in a Gray Whale Using Ultrasonography.
Michael P. Curran and William M. Asher
20 Surgical Attachment of a Telemetry Device to the Dorsal Ridge of a Yearling
California Gray Whale. Eschiichtius lubiisiiis. John C. Sweeney and Joel
L. Mattsson
22 Some Hematologic Observations on the California Gray Whale. Alfred
Zettner
24 Some Coagulation Factors in Plasma from a California Gray Whale.
Em hrii hiiiis nihii.\tii\. W. Medway
25 Fluorescent Karyotype of the California Gray Whale, Deborah A. Duffield
28 Some Physiological Parameters of the Blood of the California Gray Whale,
William G. Gilmartin. Richard W. Pierce, and George A. Antonelis. Jr.
31 Feeding of a Captive Gray Whale, G. Carleton Ray and William E. Schevill
38 Sounds Produced by the Gray Whale. Eschiichtius nihiisiiis. James F. Fish.
James L. Sumich. and George L. Lingle
45 Aerial Observations of Migrating Gray Whales, Esclirichiitis rohiisiiis. off
Southern California, 1969-72, J. S. Leatherwood
50 A Note on Gray Whale Behavioral Interactions with Other Marine Mam-
mals. J. S. Leatherwood
51 Aerial Observations of Gray Whales During 197.^. Paul N. Sund and John
L. O'Connor
52 Telemetering of Temperature and Depth Data from a Free Ranging Year-
ling California Gray Whale, Eschrichiiiis rohtisms. W. E. Evans
58 Capture and Harnessing of Young California Gray Whales. Eschrichiiiis
rohiisiKs. Kenneth S. Norris and Roger L. Gentry
Cover. — Alter almosi a year in capllvlly, the
Calilornia gray whale. Gigi II, Is carried in a
special cradle aboard ship to the site where
she will later be released. Photograph, courtesy
of Audio Visual Prod. Div., Naval Undersea
Center, San Diego. Calif.
U.S. DEPARTMENT OF COMMERCE
Frederick B. Dent, Secretary
NATIONAL OCEANIC AND
ATMOSPHERIC ADMINISTRATION
Robert M. White, Administrator
National Marine Fisheries Service
Robert W. Schoning, Director
Address correspondence to: Marine
Fisheries Review, NMFS Scientific
Publications Staff, Room 450, 1107
N.E. 45th St., Seattle, WA 98105.
Publication of material from sources
outside the Service is not an endorse-
ment. The Service is not responsible
for the accuracy of facts, views, or
opinions of these sources.
Although the contents have not been
copyrighted and may be reprinted
freely, reference to source is
appreciated.
The Secretary of Commerce has de-
termined that the publication of this
periodical is necessary in the trans-
action of public business required by
law of this Department. Use of funds
for printing this periodical has been
approved by the Director, Office
of Management and Budget, May 10,
1973.
Editor; Thomas A. Manar
Managing Editor; Willis L. Hobart
For sale by the Superintendent of
Documents, U.S. Government Print-
ing Office, Washington, DC 20402.
Price $1.25 (single copy). Subscrip-
tion price; $12.50 a year, $15.75 a
year for foreign mailing.
Marine Fisheries Review
Vol.36, No. 4
April 1974
CONTENTS
Articles
1 Preface. W. E. Evans
2 Introductory Remarks. Brian J. Rothschild
3 Respiration and Metabolism in Two Baleen Whale Calves, Eric A. Wahren-
brock. Gary F. Maruschak. Robert Eisner, and David W. Kenney
9 Ballistocardiography as a Technique for Comparative Physiology. N. Ty
Smith and Eric A. Wahrenbrock
15 Investigation of Blubber Thickness in a Gray Whale Using Utrasonography,
Michael P. Curran and William M. Asher
20 Surgical Attachment of a Telemetry Device to the Dorsal Ridge of a Yearling
California Gray Whale. Eschiichliiis rohustus. John C. Sweeney and Joel
L, Mattsson
22 Some Hematologic Observations on the California Gray Whale. Alfred
Zettner
24 Some Coagulation Factors in Plasma from a California Gray Whale.
Escliricli litis rohiisnis, W. Medway
25 Fluorescent Karyotype of the California Gray Whale. Deborah A. DufHeld
28 Some Physiological Parameters of the Blood of the California Gray Whale.
William G. Gilmartin. Richard W. Pierce, and George A. Antonelis. Jr.
31 Feeding of a Captive Gray Whale. G. Carleton Ray and William E. Schevill
38 Sounds Produced by the Gray Whale. Eschrichtiiis rohiislus, James F. Fish.
James L. Sumich. and George L. Lingle
45 Aerial Observations of Migrating Gray Whales, Esclirichtius rahiisius. off
.Southern California, 1969-72, J. S. Leatherwood
50 A Note on Gray Whale Behavioral Interactions with Other Marine Mam-
mals, J. S. Leatherwood
51 Aerial Observations of Gray Whales During 1973. Paul N. Sund and John
L. O'Connor
52 Telemetering of Temperature and Depth Data from a Free Ranging Year-
ling California Gray Whale. Escliriclilius rohusnis, W. E. Evans
58 Capture and Harnessing of Young California Gray Whales, E\i.hiii.litiiis
nihusiiis. Kenneth S. Norris and Roger L. Gentry
Cover. — Alter almost a year in captivity, ttie
Caiitornia gray whale. Gigi II. is carried in a
special cradle aboard ship to the site where
she will later be released. Photograph, courtesy
of Audio Visual Prod. Div., Naval Undersea
Center, San Olego, Calif,
U.S. DEPARTMENT OF COMMERCE
Fretlerick B. Dent, Secretary
NATIONAL OCEANIC AND
ATMOSPHERIC ADMINISTRATION
Robert M. White, Administrator
National Marine Fisheries Service
Robert W. Schoning, Director
,„o»;w^,.
Address correspondence to: Marine
Fisheries Review, NMFS Scientific
Publications Staff, Room 450, 1107
N.E. 45th St., Seattle, WA 98105.
Publication of material from sources
outside the Service is not an endorse-
ment. The Service is not responsible
for the accuracy of facts, views, or
opinions of these sources.
Although the contents have not been
copyrighted and may be reprinted
freely, reference to source is
appreciated.
The Secretary of Commerce has de-
termined that the publication of this
periodical is necessary in the trans-
action of public business required by
law of this Department. Use of funds
for printing this periodical has been
approved by the Director, Office
of Management and Budget, May 10,
1973.
Editor: Thomas A, Manar
Managing Editor: Willis L. Hobart
For sale by the Superintendent of
Documents, U.S. Government Print-
ing Office, Washington, DC 20402.
Price $1.25 (single copy). Subscrip-
tion price: $12,50 a year, $15.75 a
year for foreign mailing.
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The California
Gray Whale
Papers presented at the
California Gray Wtiale Workshop,
University of California,
San Diego, Scripps Institution
of Oceanography,
21-22 August 1972.
Sponsored by
Southwest Fisheries Center,
La Jolla Laboratory, National
Marine Fisheries Service, NOAA
and
U.S. Department of the Navy's
Naval Undersea Center,
San Diego.
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'•P
CARL L. HUBBS, Chairman.
W. E. EVANS, Editor.
PREFACE
In early March 1971 an expedition
sponsored by Sea World. Inc. of .San
Diego, under the direction of David
W. Kenney with scientific support
from the University of California.
San Diego, captured a newborn female
California gray whale [Eschrichiius
ri>hu.'itns) in Scammon's Lagoon. Baja
California Sur. Mexico. This whale,
5.84 meters long and weighing 1.952
kilograms, arrived at Sea World in
San Diego on 17 March 1971.
Although not the first successful
capture of an immature California
gray whale, this was however the be-
ginning of a successful year of mainte-
nance in captivity and the subsequent
release into the wild, the first time for
any species of baleen whale. The
results of the scientific studies con-
ducted during this year of captivity
and the later field observations which
were stimulated by the release of this
unique whale, are the subject of this
publication.
Many important contributors to
our overall understanding of the com-
plexities of the biology of the Cali-
fornia gray whale are not formally
represented in this report as contribu-
tors. The impact of the work of Carl
L. Hubbs. Scripps Institution of
Oceanography. University of Califor-
nia, San Diego, La Jolla; Raymond
Gilmore, Museum of Natural History,
San Diego; and Dale Rice, National
Marine Fisheries Service. Northwest
1
Fisheries Center. Seattle. Wash., as
pioneers in estabUshing the basis for
our present knowledge of the status
of the gray whale population cannot
be overstated.
Special acknowledgement is also
due David W. Kenney. of Poway,
Calif, for his efforts in successfully
capturing and maintaining in good
health the immature gray whale
named Gig! II. the subject of most
of the research reported here. Dr.
Kenney should be applauded for his
persistence in overcoming seemingly
insurmountable opposition. Many of
Dr. Kenney 's colleagues were doubt-
ful that a newly born gray whale
could be successfully maintained alive
in captivity for more than a few
months, let alone one year. Yet, this
INTRODUCTORY REMARKS
BRIAN J. ROTHSCHILD
It is a great pleasure to welcome
you to the California Gray Whale
Workshop. The Workshop is being
held in a significant location and at
a particularly appropriate time.
The location. La Jolla. is of course,
quite near the area surveyed as part
of the well-known California gray
whale census and is also a focal region
for other studies on the dynamics
and life history of the California gray
whale. Some of this research will be
presented at this Workshop where you
will hear about such diverse topics as
husbandry, respiration and metabolism,
cardiovascular physiology and blood
studies and behavior and physiology —
all related to the California gray
whale.
In addition to being a region where
many contemporary studies on the
gray whale have been undertaken, it
was also in this general area of the
North American coast that Charles
M. Scammon. whaler and sometime
Brian J. Rothschild is Director,
Southwest Fisheries Center, Na-
tional Marine Fisheries Service,
NOAA, La Jolla, CA 92037.
goal was achieved with overwhelming
success. From predictions of normal
growth. Gigi should have reached a
total weight of 5.946 kilograms and
a length of 8.30 meters by 20 March
1972. During her last week in captiv-
ity (6-13 March 1972). Gigi II was
weighed three or four times. Due to
the use of three different scales and
two different conditions of weigh-
ing (i.e., animal fasting and animal
fed prior to weighing), her final
weights ranged from 5,364 kg to
6,350 kg. This weight range remark-
ably brackets the predicted weight
previously mentioned. Her final over-
all length on 13 March 1972 was
8.15 meters, also significantly close
to the predicted length of 8.30 meters
based on normal growth. W.E.E.
captain in the U.S. Revenue Marine,
undertook his early studies of the
natural history of the gray whale. His
studies "The Marine Mammals of the
North-Western Coast of North Amer-
ica." were published in 1874. Many
of Captain Scammon's observations
on the gray whale were made in the
mid-1850's when he discovered a
major nursery ground of the Califor-
nia gray whale in a Baja California
embayment. Laguna Ojo de Liebre.
now frequently called Scammon's La-
goon. Scammon was also involved in
(he early, intensive harvest of this
species, an activity that was terminat-
ed in 1946 when the International
Whaling Commission declared the
gray whale a protected species.
The timing of this symposium is
also appropriate. There is now an
unprecedented interest in marine mam-
mals. TV. radio, motion pictures,
newspapers and magazines have all
contributed to a growing public aware-
ness and concern with these fascinat-
ing animals. Unfortunately, this de-
luge of publicity has resulted in a
mixture of fact and fiction. The tic-
tion has been further fed b\ various
indiscriminate interpretations which
often accompany events of high pub-
licity value, inadequate data collec-
tion, and difficulties in interpreting
the sparse marine mammal data. Fur-
ther complications arise from conflict-
ing and contradictory views of special
interest groups that influence resource
decisions. A case in point is. of course,
the blue whale.
Because this is also a time when
significant policy and conservation
decisions are being made on marine
mammals, it is particularly important
to concentrate on the generation of
factual information. The conservation
of our resources is essentially a deci-
sion-making process; this process can
only be effective if decision-makers
are supplied with appropriate facts.
Workshops such as this California
Gray Whale Workshop will do much
to contribute to our understanding
and knowledge of marine mammals
and assist in making better resource
decisions which hopefully will preserve
these Leviathans for the education
and enjoyment of future generations.
I think Herman Melville had a
premonition that all of this would
come to pass; that status of marine
mammal stocks would be of world
concern and as a small part of this
concern we would be holding our
workshop. In fact he could be before
you now saying, as he did in Mohy
Dick:
"Already we are boldly launched
upon the deep; but soon we shall
be lost in its unshored, harborless
immensities. Ere that come to pass;
ere the Pequod's weedy hull rolls
side by side with the barnacled
hulls of the Leviathan; at the outset
it is but well to attend to a matter
almost indispensable to a thorough
appreciative understanding of the
more special leviathanic revela-
tions and allusions of all sorts
which are to follow.
It is some systematized exhibi-
tion of the whale in his broad
genera, that I would now fain put
before you. Yet is it no easy task.
The classification of the constitu-
ents of a chaos, nothing less is here
essayed. Listen to what the best and
latest authorities have laid down . . ."
MFR PAPER 1045
Respiration and Metabolism in
Two Baleen Whale Calves
F.RIC A. WAHRENBROCK, GARY F. MARUSCHAK,
ROBERT ELSNER, and DAVID W. KENNEY
ABSTRACT
Eric A. Wahrenbrock and Gary
F. Maruschak are members of
the staff of the Anesthesia Labo-
ratory of the School of Medicine,
University of California, San
Diego. La Jolla, CA 92037.
Robert Eisner is a member of
the staff of the Physiological
Research Laboratory of Scripps
Institution of Oceanography.
L'niversitv of California. San
Diego, P^O. Box 109. La Jolla,
CA 92037. David W. Kenney
was formerly a member of the
staff of Sea World. Inc.. San
Diego. Calif.; his present address
is 14220 Poway Rd.. Poway,
CA 92064.
W-V perfonucd rcspiiiilnry unci mcuiholic sliulics on two female gray whale
calves. Allhongh one died 2 inonlhs after capture, the other thrived during a
year'.f captivity, pennitting serial observations while growing, and weiglied in
e.Kcess of 6.350 kg when released. They appeared to he of normal size and
weight compared to whtdes in the wild. Relative increases in body length and
weight, lung volume, minute ventilation, and metabolic rale were similar to
those in terrestrial mammals, as was the growth efficiency. Lung volume and
metabolic rate could be predicted with only partial success from the relation-
ships of those variables to body weight proposed by Tenney and Kleiber, per-
haps due to intniuturity in the whales.
Compared to terrestrial mammals, tite ratio of tidal volume to resting lung
volume in the wliule was large, while the ratio of wasted ventilation to tidal
volume was small. We measured respiratory excursions of arterial Oi and CO2
tensions of 36 and 16 mm Hg, respectively, consonant with the relationships
between respiratory rate, lung volume, tidal volume, and metabolic rate.
INTRODUCTION
Although the physiology of toothed
whales. particularU porpoises, has
been studied at some length (Irving.
Scholander, and Grinnell. 1941: Olsen.
Eisner. Hale, and Kenney, 1969;
Olsen. Hale, and Eisner. 1969; Scho-
lander. 1940), the study of living
baleen whales has been particularly
elusive. The size and dietary habits
of these large mammals present for-
midable obstacles to their maintenance
in captivity, and these obstacles are
compounded by ignorance of the
whales' growth rate, dietary require-
ments, metabolism, and hematologic
and cardiorespiratory physiology.
However, these and other aspects
of the biology of baleen whales are,
in many respects, unique among mam-
mals: research would therefore be
doubly rewarding. This line of rea-
soning led to the capture and study
of the two animals reported herein,
and to this workshop.
We were naturally inclined toward
studies of especial personal interest,
and recognize their limited scope
and serious omissions (cardiac out-
put, for example). We are here re-
porting observations on growth, res-
piratory function, and metabolic
rate: additional reports of detailed
nutritional, metabolic, biochemical
and hemotologic studies; inert and
anesthetic gas uptake; and respira-
tory mechanics will follow.
METHODS
The first gray whale calf (Gigi 1)
was captured in Scammon's Lagoon,
Baja California, Mexico in February
1965 and brought to San Diego,
where a number of respiratory and
metabolic studies were performed.
Although the whale at first seemed
to thrive, it died of an uncontrollable
infection about 2 months after it
was captured.
The second calf. Gigi II, was cap-
tured in March 1971, again in Scam-
mon's Lagoon, and was again kept
in (increasingly larger) pools at an
oceanarium in San Diego. Gigi 11
thrived indeed: gained in size, was
weaned, was studied intensively, and
was reluctantly (but inevitably) re-
leased almost exactly a year after
her capture.
Two of the authors were members
of each of the expeditions (DWK. and
RE on the first, and DWK and EAW
on the second) and one of us (DWK)
was responsible for the medical care
of both animals while in captivity.
For most of the studies reported
here, the water level in the tank was
lowered so as to nearly immobilize
the whale, leaving about 12 inches
of dorsal body surface above the wa-
ter level, and the blowhole barely
awash. A few of the studies were per-
formed with the whale completely
stranded on the bottom of the empty
tank.
The respiratory pattern in whales
and other marine mammals consists
of an expiration followed by an im-
mediate inspiration, followed by an
interrespiratory pause during which
the airway is closed. The duration of
the pause in Gigi 11 was ahout I
minute, and inspiration and expiration
together required about 2 seconds.
Two observations can be made from
this respiratory pattern (commonly
called ■■apncustic'"): I) a valve would
be needed in order to separate inspira-
tion from expiration, and 2) resting
lung volume is different from that in
terrestrial mammals, because in ceta-
ceans it includes the tidal volume.
Accordingly, we fabricated nonrc-
breathing valves: first of approximate-
ly 5 inches diameter, and later (for
Gigi ID of 8 inch stovepipe (Figure
I), thus permitting us to collect un-
contaminated exhaled gas. For Gigi 1
a large, calibrated, counterbalanced,
bellows-tvpe spirometer was used: and
for Gigi 11. expired gas was collected
in 900 liter meteorological balloons.
The volume of exhaled gas was then
measured by emptying the balloons
through a calibrated dry gas meter
(Wright Respirometer or American
Meter Co.)' at a constant, known
flowrate. Aliquots of mixed expired
gas from Gigi I were analyzed for Og
and CO2 with a .Scholander apparatus,
and for Gigi II with a moditied Hal-
dane apparatus (Lloyd-Gallenkamp).
as were samples of end-tidal gas, ob-
tained from a port just beyond the
expiratory valve leaflet of the nonre-
breathing device.
Resting lung volume was measured
in Gigi II by injecting 1.50 liters of
pure helium into the inspiratory port
of the nonrebreathing valve during
inspiration. The subsequent expira-
tion was captured, and mixed expired
gas analyzed for helium with a sensi-
tive, calibrated katharometer (W. F.
Collins).
Arterial blood was drawn Irom
Gigi II bs percutaneous puncture of
the digital artery in a flipper with an
18 gauge 3 inch needle, and arterial
placement ensured by observing pul-
sations of blood through the needle.
Because of the configuration of the
' Reference to name of firm does not imply
endorsement by tfie National Marine Fisheries
Services, NCAA
arterial and \enular system, it is pos-
sible that arterial blood was contam-
inated at times with venous blood.
Some of the gas samples from Gigi 11.
and all of her blood samples, were
analyzed for O2 and CO., tensions
(/'02 and ^002' w''h a hlood gas
analyzer (Radiometer BMS-3). with
which blood pH could also be deter-
mined.
From timed gas collections during
which the number of breaths was
also counted, respiratory rate, minute
ventilation, tidal volume, oxygen con-
sumption, and wasted ventilation (or
•dead space" fraction. ^jIV,) could
thus be determined by suitable analysis.
RESULTS
In the first few weeks of captivity,
each whale lost weight, but gained
thereafter (Figure 2). The rate of gain
during the first 8 months was about
200 kg/mo in Gigi II.2 She was
weaned at about 7 months of age. as
are calves in the wild (Rice and Wol-
man. 1971). At age 10 months, she
entered a very rapid growth phase
during which her food intake increased
from about 1.200 to about 1.800
pounds of squid/day. and her rate
of gain in weight increased almost 5
fold, to 970 kg/mo or (for those of
us who enjoy such reductions) approx-
imately -3 pounds/hour. Each whale
gained in length regularly: although
Gigi 1 was smaller than Gigi II when
captured, their increases in body size
were similar (Figure .^). This suggests
that the infection did not seriously
impede her growth.
Respiratory rate (/) was counted on
manv occasions; it varied with the
whales' activity. It averaged 2/niin
for Gigi 1 at first, and increased to 4
or .s/min after age 2 months. However,
this whale had atelectasis and pneu-
monia secondary to a harpoon wound.
2 Both calves were first fed by gavage. and in
botti trie liquid diet was gradually changed
from mainly whipping cream to a mixture of
ground squid, ground bonita, calcium casemate,
yeast, and corn oil For Gigi-ll. the proportion
of squid in the diet was gradually increased
until the lime of weaning
Figure 1. — NonbreathJng valve tor Gigi II,
constructed of 8 inch stovepipe and containing
one quarter inch neoprene foam rubber valve
leaflets. Inspiration was from the side-arm, and
the inspiratory valve leaf and its supporting ring
were slanted so that closure was assisted by
gravity.
AGE SIZE
Figure 2. — The whale was weighed with an
industrial heavy duly scale (Dynamometer) by
lifting her from the water with a crane while
supported on a canvas and pipe slrelcher, and
subtracting the tare weight. Body length was
measured on a straight line from lips to no4ch
in fluke. Dala from Gigi I are represented by
open circtes and squares, and tor Gigi ft by
solid symbols. Length in meters on left hand
scale; weight in thousands of kilograms on
right hand scale.
leaving the ineamng of this observa-
tion somewhat uncertain. When Gigi
II was motionless, or nearly so, /'
averaged 1/min, irrespective of age.
Accordingly, tidal volume ( F,) and
minute ventilation were nearly equal
(Figures 4, 5). Each value for F, is an
average of three or more measure-
ments, as we observed that \\ varied
7
6
WEIGHT LENGTH
/ 1-::
5
/
4
'■5 , /
3
2
^^^
1-
METERS
Figure 3. — Weight plotted as a
(unction of length in the two
gray whale calves. Data from
Gigi I is represented by squares,
and from Gigi II by circles. At
the time of the rapid weight
increase Gigi II was approxi-
mately 9.5 months old. Gil more
(1961) reported data from one
calf which died after being
stranded in San Francisco Bay.
Data from Rice and Wolman
(1971) represent north and
southbound calves (lower and
upper points respectively): the
difference supports the hypothe-
sis that gray whales fast during
the southern migration.
LUNG
VOLUME
.
400
/
IITE»S
/
300-
/
• /
200
/
/•
100
•
WEIGHT
kg X 10^
-[
200 -
TIDAL VOLUME
/
150-
/
-
LITERS /
100-
50-
••/
WEIGHT
1 \ 1 1—
kg X 10'
1 1 1
Figure 4. — Tidal volume (V|) in a gray whale
calf during the first year of life. Tlie regression
equation for tlie line is: V( = (47 ^ body
weigtit in metric Ions) - 70. Tlie correlation
coefficient r = 0.99.
MINUTE VENTILATION
I 4- LITERS
markedly from one breath to the next,
sometimes by 50 percent.
Resting lung volume. (Figure 6)
necessarily varied from breath to
breath also, and in addition, the mea-
surement was technically difficult be-
cause of the difference between in-
spired and expired \\. Nevertheless,
five measurements were felt to be
adequate. Two measurements were
made at weight = 6.150 kg. one
when awash and one when stranded
on the bottom of the completely
drained tank. Lung volume was re-
duced by about 20 percent by strand-
ing, although that value was deter-
mined only once.
From time collections of mixed ex-
pired gas. oxygen consumption was
computed (Figure 7). The composi-
tion of end tidal gas from Gigi I!
was determined on several occasions,
and did not vary systematically with
age. End tidal ^02 varied from 54
Figure 5. — Minute ventilation
(Vf) in two gray wtiale calves,
expressed as a function of body
weigfil. Ttie triangle represents
data from observations in Gigi I.
Ttie regression equation for Itie
data from Gigi II (circles) is:
V£ = (70 X body weight in
metric tons) - 117. Although
the r = 0.94 for this rectilinear
regression, it is apparent that a
sigmoid curve could be even
more closely filled to the data.
Figure 6. — Resting lung volume in a gray whale
calf (Gigi II), expressed as a function of body
weight. Resting lung volume includes tidal
volume, and is the volume of gas in the lungs
during the intervals between breaths. The
equation for the line is: lung volume = (70 X
body weight in metric tons) - 44; for which
r = 0.94.
Table 1. — Arterial blood gas tensions and pH,
drawn at random during the respiratory cycle.
Age, months
2 5 3 0 10
55 62 60
56 69 41
7 23 7,32 7 35
Pa02, mm Hg
PaC02, mm Hg
pH
OXYGEN CONSUMPTION
WEIGHT kg X 10^
WEIGHT kg X lO'
H 1 1 H
3 4 5 6
Figure 7, — Oxygen consumption in liters/min
(Vq.,) in two gray whale calves, expressed as
a function of body weight. The triangle rep-
resents data from observations in Gigi I. The
regression equation tor the data from Gigi II
(circles) is: VOj = (41 « body weight in
metric tons) - 5.7; lor which r = 0.96.
70
-Po,=
3 6 lorr
60-
. ^
'^ Pco, =
16 torr
50-
^
^^^
40-
^^^
'
30-
. Po
CO2
20-
10-
■
Poo,
1 1 1
Figure 8. — Arterial O2
and Cq., tensions (Pao^
and PacOj) '" ^ 9"*
wtiale cal'f, from se-
quential blood samples
drawn about every 15
sees during five respira-
tory cycles.
so 60 70
to 85 mm Hg, and the corresponding
Pcoo varied from 75 to 54 mm Hg.
We computed wasted ventilation from
the difference between end tidal and
mixed expired Pco2- " equalled '3.0
percent of V^ at age 3 months and
13.5 percent at age 13 months.
Arterial Po2' ^C02- ^^^ P^* ^'^■'^
measured on three occasions in sam-
ples drawn at random during the res-
piratory cycle. (Table I). Those values
also varied considerably: the differ-
ences between arterial and alveolar
Pq„ and Pcoo ^^"^^ difficult to inter-
pret, and were sometimes negative.
Therefore, we measured blood gases
and pH in arterial blood drawn se-
quentially during the respiratory cy-
cle (about every 15 sec); the values
then varied systematically (Figure 8).
DISCUSSION
As we are presenting data concern-
ing the respiratory and metabolic
changes in growing whales, we should
examine the hypothesis that their size
and rate of growth were normal.
There is ample reason to raise the
question, for confined animals fed
contrived diets should always be sus-
pected of exhibiting biological values
which would be abnormal for the
population in nature. There are two
methods of examining this question:
to make comparisons with other gray
whales; and to look for internal evi-
dence of abnormal growth and de-
velopment.
Data from Gilmore (1961) and
Rice and Wolman (1971) comprise
the first method, for they have exam-
ined gray whale calves similar in age
and size to Gigi II. Her weight and
length at the time of release compare
favorably to the other data on calves
thought to be yearlings (Figure 3).
As gray whales in the wild are thought
to fast during the southern migration
(Rice and Wolman, 1971), the obser-
vation that Gigi II was slightly heavy
for her length should be interpreted
with caution.
The internal evidence relating to
the question consists of the observa-
tion that Gigi II sustained an increase
in body length which preceded any
considerable growth in body weight,
mitigating against an argument that
she was grossly overweight or overfed,
and which is consistent with the pat-
tern of early growth in other mammals
(i.e.. exponential for weight and linear
for length) (Christian, 1972; Carlan-
der and Ricker, 1962; Brody, 1964).
Although there is some disagree-
ment (Gilmore. 196 l.andpers. comm.).
newborn gray whales are estimated to
be 4.9 meters in length at birth (Rice
and Wolman. 1971). Their birth-
weight is less certain, although Rice
proposes the weight of the products
of conception at term to be between
1.000 and 2.000 kg. Our estimates
of the birthweight of the two whales
at about 1.500 kg. and the birthlength
at just under 5 meters are therefore
consistent, and permit extrapolation
of their ages at capture to 4 weeks for
Gigi I and 10 weeks for Gigi II. At
age 1 year. Gigi II had increased her
birthweight by about 3.5 fold, and
her birthlength by a little less than
1.5 fold.
Comparison of growth rates be-
tween species may be deceptive due
to species differences in longevity,
newborn maturity, and adult body
size. However, since humans and
gray whales (Rice and Wolman. 1971)
have similar life spans, and are both
large mammals, it is of interest to
consider their relative growth rates
in body weight, respiration, and me-
tabolism.
If mature female gray whales are
13 meters in length (Rice and Wol-
man. 1971; Scammon. 1874). and 30
to 35 thousand kg in weight, then the
newborn whale must increase its
birthweight by about 20 fold and its
birthlength by about 2.5 fold. The
fractional annual and ultimate in-
creases in weight of the whales are
similar to those in man. but the frac-
tional increase in length is greater
(Benedict and Talbot. 1921).
Lung volume in human infants in-
creases as a cubic function of body
length (Cook and Hamann. 1961). a
satisfying observation considering the
geometry involved. The regression
onto body weight is not very reliable;
the regression onto length is approxi-
mately; Total Lung Capacity (liters)
= body length (meters) cubed. Al-
though we have insufficient data for
the whale to make firm conclusions
(and have arbitrarily linearized the
data against body weight in Figure 6).
calculation of resting lung volume
against body length suggests that an
equation of Lung Volume = 0.62 X
length' also fits the data. This rela-
tionship seems reasonable, as resting
lung volume in the whale is probably
less than total lung capacity.
Tidal volume and minute ventila-
tion increased during the year's growth,
and as a first approximation, we have
again linearized the data (Figures 4.5).
We recognize that changes in res-
piratory function and metabolic rate
are probably not rectilinear functions
of body weight, but we also recognize
that the data available to us represent
only a small portion of the full curves.
Tidal volume increased about 8 fold
as body weight tripled: the correspond-
ing value for minute ventilation was
10 or 12 fold. Comparison of this
growth rate with that of terrestrial
mammals is awkward, because of
marked changes in their respiratory
rate during growth (Watson and Low-
rey. 1962). while respiratory rate in
the whale was constant. In terrestrial
mammals, tidal volume changes as a
function of lung volume (which in
turn varies as a cubic function of
length), while minute ventilation
changes as a function of metabolic
rate (Tenney and Kemmers. 1963);
a more complex function of growth,
which is not linear on any conve-
nient parameter of body size because
of growth spurts during early and
late childhood (Benedict and Talbot.
1921).
The increase in the metabolic rate
of Gigi II corresponded to the in-
crease in her ventilation, and for a
tripling of weight, increased about 10
fold. This increase is of the same order
as the increase in human metabolic
rate during the first year (about 8
fold) (Benedict and Talbot. 1921).
and is consistent with our general
impression that growth in the whale,
whether of body weight, lung volume,
or metabolic rate, proceeded in paral-
lel with, or only slightly more rapidly
than, human growth.
Observation of animals at the ex-
tremes of body size invites inter-
species comparisons of biological
phenomena. The existing data for two
such correlations; of lung volume
with body size (Tenne\ and Remmers.
1963), and of metabolic rate with
body size (Kleiber. 1961), are par-
ticularly well organized. Tenney has
shown that lung volume is closely
related to body mass (over a range
of body mass of 5 orders of magni-
tude) by the equation: log lung vol =
1.02 log body weight — 1.25, with
volume in liters and weight in kg.
This yields a predicted lung volume
in Gigi II of 410 liters at 6,150 kg,
which corresponds closely to our
measurement of 428 liters, but which
diverges widely from values obtained
early in her growth. Tenney measured
total lung capacity (TLC) of excised
lungs, and we measured resting lung
volume: this ordinarily considerable
difference is fortuitously minimized
by the fact that resting lung volume
in the whale is a larger fraction of
TLC than is the case for terrestrial
mammals. This measurement permits
considerable extension of Tenney's
data, for his largest animal, also a
cetacean, weighed only 1,750 kg.
Kleiber studied the metabolic rates
of animals also differing in body size
by about 5 orders of magnitude, and
concluded that metabolic rate was
best related to the 0.75 power of
weight, by the equation: log A/ = 1.83
+ (0.756 log W) ±0.05, with M in
kcal/day and W in kg. Using the con-
version factor of 4.8 kcal = 1 liter
of O), the whales" metabolic rates
compare favorably with that regres-
sion line up to a body weight of 3,000
kg. but diverge significantly there-
after. The last metabolic rate mea-
sured was 16.8 l/min. while the cal-
culated value from Kleiber's equation
is 6.8 l/min. It is notable that metab-
olism in Gigi II. Benedicts elephant,
and Irving's whale all differ from
Kleiber's prediction, thereby raising
the question of whether large mam-
mals do indeed follow the 0.75 power
rule. However, the value for the
70.000 kg fin whale was extrapolated
from a measurement in a porpoise
(Irving. Scholander. and Grinnell.
1941). and neither the elephant nor
our whales were studied under condi-
tions meeting Kleiber's criteria of
ambient temperature neutrality, adult-
hood, and basal postabsortive state.
The resulting errors would be in the
direction of the observed differences.
Divergence from the 0.75 power rule
may also be seen in growing cattle,
horses, children, and rodents (Brody.
1964).
During the phase of rapid weight
gain. Gigi II ate from about 1.200 to
about 1.800 pounds of squid per day,
and gained weight at the rate of
about 980 kg/mo. If we assume thai
squid are about 80 percent water
and the dry weight is equivalent to 5
kcal/gm (R. Lasker. pers. comm.).
and make the further assumption that
growing whale tissue contains the
same energy (1.720 kcal/kg wet
weight) as other growing mammalian
tissue (Mayer. 1949), it is possible
to calculate the gross efficiencies for
growth of a baleen whale calf of 10.3
percent and 6.9 percent. Correcting
for metabolic rates of 11.0 and 16.8
liters O'/min yields net efficiencies for
growth of 12.0 percent and 8.0 per-
cent (Brody. 1964). In general, growth
efficiency is independent of body size
(Kleiber. 1947), but is a diminishing
function of metabolic age: the calculat-
ed values are within the expected
range for terrestrial mammals beyond
the first doubling of body weight
(Brody, 1964).
Tidal volume equalled about 50
percent of resting lung volume, ir-
respective of age or size. This is a
smaller ratio than that reported for
other diving mammals (Irving et al..
1941; Olsen et al.. 1969; Scholander,
1940), although they were mature.
The ratio of wasted ventilation to
tidal volume (VjjVj) in Gigi II was
about 13 percent, irrespective of age.
This value is consistent with observa-
tions in mature diving mammals (Ir-
ving et al., 1941; Scholander. 1940.
and Kooyman. pers. comm.). and is
considerably smaller than the ratio in
terrestrial mammals. However. I'j/
Vj diminishes with increasing I'^ in
humans and dogs (Bouhys. 1964). a
pertinent observation in view of the
relatively large V^ in the divers.
Fluctuations in arterial Pq and
A:02 *''h respiration have been pre-
dicted in man and terrestrial animals
(Otis. 1964; Suwa and Bendizen. 1972)
and diving animals (Irving et al.. 194 I).
Those fluctuations are influenced by:
1) the relative sizes of the tidal and
resting lung volumes; 2) the relation-
ship between resting lung volume and
metabolic rate; 3) the relationship
between the fluctuating pulmonary
blood flow and the fluctuating alveolar
gas composition (Otis, 1964): and 4)
the solubility of respiratory gases in
pulmonary tissue. The greatest differ-
ences we observed were A/'og = 36
mm Hg and A/coo ~ '^ "^^n Hg
(Figure 8): These considerable excur-
sions follow from the ratio of resting
lung volume to tidal volume, which
in the whale is about 2 and in man
about 5; the ratio of resting lung
volume to metabolic rate, which in
the whale is about 20 and in man is
about 10; and the very large differ-
ence between human and whale res-
piratory rates. Taken together, these
relationships suggest that apneustic
breathing in the whale is just as it
seems: each breath interrupts a res-
piratory pause which actually repre-
sents a period of breathholding, dur-
ing which appropriate changes occur
in arterial blood gas tensions. Al-
though tempting, approaches toward
cardiac output computation are ham-
pered by ignorance of the composition
of mixed venous blood. Calculations
of the "mean" alveolar gas or arterial
blood composition are similarly ham-
pered, and by imprecision in the
sample collection timing as well.
CONCLUSIONS
1. These two gray whale calves
have provided the first opportunity
for the collection of physiologic data
from living baleen whales. The growth-
rate in one of them was such that she
became the world's largest captive
animal.
2. Comparison of their size with
that of whales in nature, and of their
growthrate with one another and with
other animals, strongly suggests that
their size and growthrate were normal.
3. We observed increases in res-
piratory function and metabolism
during growth similar to the increases
in terrestrial mammals. In particular:
relative increases in body weight: and
of lung volume, minute ventilation,
and metabolic rate as functions of
body weight, proceeded in approxi-
mate parallel to the relative increases
observed in man,
4. Interspecies comparisons of ab-
solute lung volume and metabolic
rate can also be based on body weight.
Where the gray whale calves differed
from correlations drawn between mam-
mals including those at the extremes
of body size, the departure could be
explained by the whales' immaturity.
5. One of the whales entered a
rapid growth phase, during which it
gained approximately 1,000 kg/mo.
Its gross efficiency for growth, cal-
culated from the amount it ate and
weighed, diminished from about 10
percent at a body weight of about
3,000 kg to about 7 percent at a body
weight of 6,350 kg.
6. The relationships between tidal
volume, resting lung volume, and
wasted ventilation are similar in the
gray whale calf to these in other div-
ing mammals: although those rela-
tionships are different from the ones
in terrestrial animals, they follow
from the apneustic respiratory pat-
tern (of infrequent but very large
breaths interrupting long periods of
breathholding at high lung volume).
7. The apneustic pattern of breath-
ing also results in respiratory excur-
sions in arterial oxygen and carbon
dioxide tensions much larger than
those predicted in terrestrial mammals.
ACKNOWLEDGMENTS
Both whales were kept in pools at
Sea World, Inc., an oceanarium in
San Diego, Calif. The staff and ad-
ministration extended to investiga-
tors courtesies and facilities both
large and small, ranging from hot
showers and coffee to underwriting
the expeditions and the subsequent
upkeep of the whales: and thereby
made these studies both possible and
enjoyable. In particular. Bud Dona-
hoo and Sue Bailey provided Gigi II
and her investigators with under-
standing and with expert assistance.
We acknowledge with thanks the sup-
port of Robert Peterson and the crew
of the FiilcDii involved in the capture.
transport, and maintenance of Gigi I.
We are also grateful for the consider-
able assistance of Jack Schultz, Ken
Hamai, James Wright. Brian D' Aoust,
and Morgan Wells. Finally, we are
grateful to the two Gigis, for they, by
patiently enduring our several insults,
made possible observations new to
the largest and least accessible sub-
order of living mammals.
LITERATURE CITED
Benedict, F. G., and F. B. Talbot. 1921.
Metabolism and growth from birth to
puberty. Carnegie Inst. Wash., Publ. No.
302, Wash., D.C.
Bouhvs, A. 1964. Respiratory dead space.
In W. O. Fenn and H. Rahn (editors).
Handbook of phvsiologv. Respiration, Sect.
3. Vol. 1, p. 699-714. Am. Physiol. Soc,
Wash., D.C.
Brody, S. 1964. Bioenergetics and growth.
Hafner, New York.
Carlander, K. D., and W. E. Ricker. 1962.
Postnatal vertebrate development. In P. L.
Altman and D. S. Dittmer (editors).
Growth, p. 33.V378. Fed. Am. Soc. Exp.
Biol., Wash., DC.
Christian, I.. L., L. N. Hazel, J. P. Scott, M.
Shelton, G. M. Sidwell. N. R. Ellis, W. W.
Swett. G. S. Templeton. and G. Van
Wagener. 1972. Growth: Mammals other
than man. In P. L. Altman and D. S. Ditt-
mer (editors). Biol, data book. p. 207-216.
Fed. Am. Soc. Exp. Biol., Bethcsda. Md.
Cook, C. D., and J. F. Hamann. 1961. Rela-
tion of lung volumes to height in healthy
persons between the ages of .S and 38
years. J. Ped. 59:710-714.
Gilmore, R. M. 1961, The story of the gray
whale. 2nd ed. Privately publ., San Diego,
16 p.
Irving, L., P. F. Scholander, and S. W. Grin-
nell. 1941. The respiration of the porpoise,
Tiirsuip.'i irunciilii.s. J. Cell. Comp. Physiol.
17:14.5-168.
Kleiber, M. 1947. Body size and metabolic
rate. Physiol. Rev. 27:5 1 1-541.
. 1961. The fire of life. An introduc-
tion to animal energetics. John Wiley &
Sons, Inc.. New York, 454 p.
Mayer, J. 1949. Gross efficiency of growth
of the rat as a simple mathematical func-
tion of lime. Yale J. Biol. Med. 21:415-419.
Olsen, C. R., R. Eisner, F. C. Hale, and
D. W. Kenney. 1969. -Blow' of the pilot
whale. Science 163:953-955.
Olsen, C. R., F. C. Hale, and R. Eisner. 1969.
Mechanics of venlilalion in the pilot whale.
Resp. Physiol. 7:137-149.
Otis, A. B. 1964. Quantitative relationships
in steady-state gas exchange. In W. O.
Fenn and H. Rahn (editors). Handbook of
physiology. Respiration. Sect. 4, Vol, 1.
p. '681-698. Am. Phvsiol. Soc, Wash., D.C.
Rice, D. W., and A. A. Wolman. 1971. The
life history and ecology of the gray whale
(Eschruhiiiis robt4snis). Am. Soc. Mammal.,
142 p.
Scammon, C. M. 1874. The marine mammals
of (he North-western coast of North
America. John H. Carmany and Co., San
Francisco, 319 p.
Scholander, P. F. 1940. Experimental inves-
tigation of the respiratory function in divid-
ing mammals and birds. Hvairad. Skr.
22:1-131.
8
Suwa, K., and H. H. Bendixen. \912. Pul-
monary gas exchange in a tidally ventilat-
ed single alveolus model. J. Appl. Phvsiol.
32:834-841.
Tenney, S. M., and J, E. Remmers. 1463.
Comparative quantitative morphology of
the mammalian lung: diffusion area.
Nature 197:54-56.
Watson, F.. H., and G. H. Lowrey. 1962.
Growth and development of children. 4th
ed.. Year Book Medical Publishers. Inc.,
Chicago, 384 p.
MFR Paper 1045. From Marine Fisheries Review, Vol.
36, No. 4. April 1974. Copies of this paper. In limited
numbers, are available from D83. Tec finical Information
Division, Environmental Science Information Center.
NCAA, Washington, DC 20235.
MFR PAPER 1046
Ballistocardiography as a Technique
for Comparative Physiology
N. TY SMITH and ERIC A. WAHRENBROCK
ABSTRACT
The ultra low-frequency haUistocarttioi^rcini was recorded on a young Califor-
nia gray whale. The tracing /.v remarkably similar to those obtained from man
and mouse, both in amplitude and in form. The IJ amplitudes fr>r mouse, man.
and whale were 2.6, 4,3, ami 4.6 cntlsec'-. We conclude that greater differences
are caused by poor recording technique or by disease than by species differences.
The major interspecies differences were seen in the timing of cardiac events,
such as preejeclion or ejection time. These differences could be caused by
differences in heart size.
The ballistocardiograph (Beg) is a
device for evaluating the mechanical
function of the heart. It has been
recorded in an incredible array of
animals, ranging from egg embryos to
cattle. One of the more interesting
facts to arise from these recordings
is that the tracings are remarkably
similar among species, particularly
mammals. This similarity holds both
in form and in amplitude. It was
therefore an excellent opportunity to
N. Ty Smith is an Associate
Professor of Anesthesia at the
University of California at San
Diego, Veterans Administration
Hospital, San Diego, CA 92161,
and Eric A. Wahrenbrock is an
Assistant Professor of Anesthesia
at the University of California,
San Diego,
extend these observations to Gigi,
an animal with an entirely different
mass and configuration from other
mammals previously used.
The Beg records the movements
of the body caused by movements of
blood in the body. First recorded in
1887. the Beg has undergone a series
of ups and downs in its attempts to
become a useful tool for measuring
cardiovascular function noninvasively.
Not until the 1950"s when physicists
and engineers entered the held, did the
Beg finally re-emerge as an accurate,
relatively simple technique.
Essentially, the Beg works on the
principle that an attempted shift in the
center of mass of a floating body is
compensated for by a movement of the
body in the opposite direction, so that
the center of mass remains constant in
relation to a fixed point. Thus, if blood
moves in one direction after ejection
by the left ventricle, the body will
move in the opposite direction. These
movements are quite small, but the
reader has certainly noticed a slight
bodily movement as he lies quietly
on a bed or a slight movement of
the pointer on a weighing scale, each
movement synchronous with the
heart beat. This minute body move-
ment can be recorded as displace-
ment, velocity, or acceleration. Figure
1 shows examples of normal tracings
in man. The important fact to note
is that the major components of the
Beg occur during ejection of blood,
particularly during the early portion.
METHODS
When the physical scientists entered
the field, they laid down certain
standards for recording the Beg,
standards which were to convert bal-
listocardiography from a haphazard
technique to a precise one. The first
requirement is that a very light bed
is necessary, in contrast to the heavy
ones formerly used. A ratio of 10:1
for subjectibed is minimal. Second,
coupling, or binding, of subject to bed
must be as tight as possible. Third.
coupling to ground must be minimal,
so that ambient vibrations can be
attenuated. The Beg is an extremely
sensitive instrument. Peak displace-
ment is about \00iJ. peak acceleration,
a few millig"s. g being the accelera-
tion of gravity. With older instruments,
vibrations from a truck outside the
building were able to destroy a bal-
Phono
EKG
Rl Hfort
listocardiographic recording. Finally,
the natural frequency of the entire
system should be as low as possible —
0.3 Hz or less is mandatory. These
four requirements imply that the ideal
Beg system is one in which subject
and bed tloat as a unit in space.
Several ingenious systems, some
simple, some complex, have been
assembled to accomplish the above
requirements. Beds have been con-
structed from aluminum and canvas,
styrofoam. balsa, or aluminum honey-
comb, and suspended by wires or
floated on mercury or air. The sim-
plest and original bed is based on the
pendulum, and was the type used in
this stud\ . The Beg bed was the same
stretcher used to weigh Gigi (Figure 2).
The stretcher was constructed from
canvas and two 20-fool heavy wall,
galvanized steel pipes 3 inches in
diameter. The total weight of 227 kg
may seem large to most ballisto-
cardiographers. but Gigi's weight at
the time was 4..'i00 kg. and the whale:
Figure 1. — Examples of normal ballistocardio-
graphic tracings in man. From top to bottom
are recorded acceleration (A), velocity (V).
and displacement (D). In addition, the EKG
and the major events of the cardiac cycle are
given as reference points. (From Scarborough
et al.. Am. J. Cardiol. 2:613-641. 1958.)
bed ratio of 20; 1 was more than
adequate. Six ropes supported the
poles, four at the ends, each 13 feet
in length, and two in the middle. A
board inserted between the two middle
ropes prevented injury to the animal.
The six ropes were suspended by a
single cable from a crane. During the
recording the cable was 7'/2 meters
from pulley to hook, giving a natural
frequency of about 0.18 Hz. The
crane was part of a truck hoist, which
was ideal for isolation from ground
because of the pneumatic lift and
the rubber tires.
Most of the water was drained from
Gigi's tank to reduce her mobility and
to enhance our own. She was reluctant
to lie on the bed. and had to be coaxed.
The coaxing process took 45 minutes.
Once on the bed. she became surpris-
ingly quiet, which was fortunate,
since she could easil\ have demolished
our fragile accelerometer. One re-
adjustment of the relative position of
whale and bed was required to level
the bed.
Acceleration was transduced in
the head-foot direction with an
Endevco' piezo-resistive accelerometer
clamped to one of the steel poles with
a large C clamp. The accelerometer
was calibrated with a pendulum, ac-
cording to the method of Moss (1961).
Lead two of the ECG was recorded
using 4 inch 18 g spinal needles. All
electrical cables were supported by a
rope stretched across the tank. A 60 Hz
passive notch filter and a .^d Hz
low pass Butterworth filter were used
on both the ECG and Beg to eliminate
unwanted noise and at the same time
preserve timing relations. Data were
recorded on a Hewlett-Packard oscil-
loscope and an Ampex FM tape rec-
order.
'Use of trade names in this publication does
not imply endorsement of commercial products
by \he National f^arine Fisheries Service.
10
RESULTS
Figure 3 shows the Beg recorded
from Gigi. In amplitude and form, it
is similar to that seen in man. Figure
4 demonstrates that the inlluence of
ventilation on the tracing is profound.
In fact, during expiration and in-
spiration reading the Beg is impossible.
Figure 5 displays the Begs of
three animals — a mouse, a man. and a
whale. Their similarities are more
striking than their differences. This
similarity holds in spite of differences
in body mass and form, amount and
distribution of fat. and natural
environment.
Table 1 lists some measurements
derived from the Bcg's of the mouse,
man. and a whale. It also gives some
fundamental values which are helpful
in comparing the species.
The Beg has been used to estimate
cardiac output and stroke volume in
several species. By using the Starr
formula (Starr and Noordergraaf.
1967. p. 177- 180) we estimated Gigi's
stroke volume to be 7.2 I. and the
cardiac output as .^08 1/min (Table I).
DISCUSSION
One of the major postulated objec-
tions to the Beg is that the amount
and distribution of body fat can con-
siderably alter the recording. This
did not seem to be the ease in Gigi.
in spite of a 3' 2 inch layer of
blubber. It is true that the old direct-
body Beg used in the 1950's was
subject to influence by body fat. How-
ever, the ultra low-frequency bed. by
virtue of its light weight and strong
coupling between subject and bed.
has eliminated most of this inaccuracy.
The fundamental natural frequency
of the body ("bowl of jelly" phenom-
enon alluded to by some in refer-
ence to the Beg) does not depend on
bod\ mass, amount of fat. or age
(Burger. Noordergraaf. and Ver-
hagen. 1953; Burger and Noorder-
graaf. 1956; Talbot and Harrison.
1955; Tannenbaum. Vessell. and
Schaek. 1956; Weissbaek. 1960a.
1960b; Tischenko. 1963). Some of
Figure 2. — Gigi. Beg bed. man. and hoist. The accelerometer is being attached to the right side
of the proximal pole. The truck was jammed against the retaining wall of the tank. A white rope
strung across the tank supports the cables.
the higher mode frequencies may
depend on the amount and distribu-
tion of body fat.
A crucial factor in ballistocardiog-
raphy is the orientation of the aorta
in relation to the body. This is so
because usually complexities have
forced ballistocardiographers to
record the Beg in one dimension, the
head-foot direction, instead of the
possible three dimensions and six
degrees of freedom. Thus if the
direction of ejection and runoff is
different in different species, the com-
parison would be difficult. The orienta-
tion of the aorta seems to be no
11
ECG -n
ULFBcq^
TIME
r^
,'* — ^
• ,o„ — ^ 1^ ^^ '~\ — ^ '~', ■ f"', — ^ f~\ — ^ r-\
J^ V/vJ^ \ . f^
^^^v^^W'-'V
Figure 3. — ECG lead two and acceleration Beg in a California gray whale. Paper speed - 50 mm/
sec. The Beg calibration of 3 cm/sec- is shown in the lower right.
ECG-n '~';—.^ ';—'!' — ^/'~^(^~^,'-^^ — """f^' — "^ r~\'"~^ f'l^ f-\ '^r-^^«*- /,-^'^~~-^'' — '^r^''~~^r~^'' — ",'"'■
ULF Bcqo
TIME
^^AJ';U\wA^wvA^\^;'vvv';V^ ■ ,^ , /. v^^v'^'"'^'-'^^
. J «v
Figure 4. — The Beg with Gigi during one breath. Paper speed = 25 mm/sec. The respiratory
influence on the Beg is considerably greater in the whale than other species. This is probably due
to the necessarily rapid and large tidal exchange.
MOUSE
An 17.25. R I9h, 27.04g . 16.8.68
,v— Av-v^/W-^^i^V^^
2cm
-lO.lmV
MAN
ULF Be
q„ A^v^/yA^~^YV-^YV-'Y^ ^
ECG-n-
WHALE
ECG-n
ULF Bcg„
TIME
-V^^ — \^^ — V-^
3cm
sec 2
Figure 5. — The ultra low-frequency acceleration Bcg's in a 27-gm mouse (top record), a 73.000-gm
man (middle record), and a 4.S00.000-gm whale (lower record). The ECG s are also shown. Note
the more rapid paper speed in the mouse Beg. Considering the 167.000-fold difference m body
mass, as well as the differences in body shape, amount and distribution of fat. and instrumentation,
the records are remarkably similar. The mouse Beg is from Juznic. G., Bibl. Cardiol. 26:281-291. 1970.
The human Beg is courtesy of Dr. Aaron G. Dinaburg.
different in the whale from other
mammals. Green (1971) describes the
course and relations thus: "Leaving the
left ventricle, the aorta makes the
characteristic left arch before passing
superficially and caudally to lie just
under the center of the thoracic
cavity to pass through the diaphragm."
Body acceleration, which is closely
related to blood acceleration, is a
constant factor among various mam-
malian species. A peak-to-peak body
acceleration of 2.5-5 cm/sec- (about
2.5-5 millig) seems to be optimum. If
the acceleration is greater, as with
severe aortic insufficiency, the slight
motion now becomes quite noticeable.
If the whale's body acceleration were
proportionateK large in relation to its
mass, the motion could become un-
comfortable. Why a smaller normal
acceleration would not be feasible, or
indeed why an initial ventricular
impulse is necessary at all. is difficult
to guess.
Other constants occur among
mammals. For example, with rare
exceptions such as the giraffe, arterial
blood pressure is very similar in differ-
ent species (Altman and Dittmer.
197 1. p. 405-4(18). One could spec-
ulate why these values are so appro-
priate. If normal arterial pressure were
higher, either the vessel walls would
have to be of considerably stiffcr
material, or the\ would have to be
so thick that the ratio of wall thick-
ness to lumen would be impractical.
If normal pressure were lower, perfu-
sion through the necessarily small
capillary vessels would be diflicult.
Perhaps even more pertinent a con-
stant involves the relative masses of
the heart and body in different mam-
mals (Table 1). Apparently there is
more variation within species than
among species.
The general form o\ Gigis Beg is
very similar to that given for normal
man h\ .Scarborough et al. ( 1958). One
can certainly recognize an HIJ ci>m-
plex and an LMN complex. It seems
that greater differences in amplitude
and lorni are caused by faulty tech-
nique, such as a heavy bed and poor
12
coupling, or by disease states, than by
differences in species. Figure 6 gives
an example of this. It compares a
virtually normal Beg in a dog with the
Beg in a dog at the terminal stages of
rejection. The latter tracing is obvious-
ly grossly abnormal and demonstrates
the extreme in Beg abnormality.
Other conditions which can cause a
greater ballistocardiographic variation
within than among species include
anginal attacks, severe coronary
artery disease. hyperthyroidism.
aortic valvular insufficiency, and
congestive heart failure (Starr and
Noordergraaf. 1967). Even a pro-
gram of physical conditioning over
several months can alter an individual's
Beg to as great an extent as the differ-
ences seen among species (Elsbach
et al., 1970, Holloszy et al.. 1964).
The major difference among the
Bcg"s of various mammals seems to be
one of timing of the systolic wave
forms. As body size increases, the on-
set of the systolic complex is delayed
(QH interval) and the complex spreads
out (HJ and HI. intervals. Table 1).
If we consider the tip of the H wave
as the onset of ejection, we shall at
worst slightly underestimate ihe
cardiac pre-ejection period. Certainly
the relative values among species can
be estimated by the QH interval. Sim-
ilarly, ejection time can be estimated
by the HL interval. This interval did
not seem to be so relatively prolonged
in Gigi as the QH. The contribution
of prolonged conduction time in
hearts of different sizes to the inter-
species differences in systolic time
intervals is probabK considerable, as
is shown b\ the PR and QRS intervals
in Table 1 .
In general, heart rate and ejection
time are inversely related. Thus part
of the differences in systolic time
intervals is due to heart rate differ-
ences. But heart rate cannot explain
all of the differences. Gigi's heart
rate of 43 beats/min was not as slow
as expected and occurred presumably
because she was excited. An athlete
with a heart rate of 40-45 beats/min
does not show the prolonged pre-
Table 1. — Some comparative values among mouse, man, and whale.
Weight (gm)
Length (cm)
Heart/body mass
(gm/100 gm)
Heart rate (beats/mm)
Beg IJ amplitude
(cm/sec-)
Beg IJ amplitude
(corrected, cm/sec^)
Bog IJ amplitude
(dynes)
Cardiac output (l/min)
Cardiac index
(Ml/mm/kg)
Stroke volume (ml)
Stroke index (ml/kg)
PR interval (msec)
QRS interval (msec)
QH interval (msec)
"Pre-eiection period"
HJ interval (msec)
HL interval (msec)
"Election time"
-t- With tail
■ Corrected for mass of bed IJ
= Measured in Gigi
I Walker, et al , 1968
- Altman and Dittmer, 1971, p. 240,
' Altman and Dittmer, 1971, p. 236-7
^ Altman and Dittmer, 1971, p. 239.
Mouse
Man
Whale
Ratio
Whale/Mouse
27
6 5-9.5 (12 5-20) + (
0 41-0 51 (■-)
73.000
) 180
0 44-0 57(3)
4,500.000
760
0 32-0 50(r)
167,000
38-117
0.78-0 98
300-700 (■•)
2.6(6)
60-80
4 3(6)
43
48
0.13
1.8
3.4-
4.7-
50-
1.5
73.0
250 X ^o■■' (;.»)
2 27X lOr
311.000
—
5.0-8 0
70-90 (9.111)
308
68.4
—
42(")
22(11)
27
70-90 (9..I")
0 9-1 2 (9.11)
180-200
80-100
90-110 ('.»)
7150
1.6
294" (320) (")
103' (90-120) (It)
320
70
12.9
11.9
43
64
140 ('.8)
320 ('.«)
205
490
4.8
7.7
/Tota!Mass\ :! '^"-"''^-" ^"t°""^^^' ^^^^' " 3"°
I p„^, ..,,, I '■ Juznic. 1970
VBody Massy . Starr and Noordergraaf, 1967.
« IVloss. 1961
9 Cullen, et al . 1970.
I" Altman and Dittmer, 1971, p. 323-
" Altman and Dittmer, 1971, p. 278.
ECG
n 4- 1 1 1 1- 1—
ULF BCGa '-wU ^"^-^-ni| Vjw.
',yv-%AA^,^,
PG
Timo. In seconds
Figure 6a. — These (wo tracings are from a conscious dog after cardiac autotransplantation. The
Beg is essentially normal. PG = Pneumogram (Whitney gauge).
ECG-n
ULF BCGa
-!^^--
^-A-.v^->V'
PG
Time in seconds
Figure 6b. — These tracings are from a dog in the terminal stages ol rejection after cardiac allo-
transplantation. The difference between the ballistocardiographic records from the two dogs is
obviously greater than that between Ihe tracings from a whale and mouse (Figure 5).
13
ejection period and ejection time that
Gigi does (Weissler et al.. I960;
Leighton et al.. 1971).
We said that the Beg is used to
estimate cardiac function. This is
possible because of the close relation-
ship between the acceleration Beg and
the acceleration of blood out of the
left ventricle into the aorta (Winter
et al.. 1966, 1967: Smith. Van Citters,
and Verdouw. 1970; Deuchar. 1966).
The latter is a proven sensitive indi-
cator of cardiac function (Noble.
Trenchard. and Guz. 1966; Noble,
Gabe. and Trenchard. 1967; Rushmer.
1964. 1970).
Gigi's stroke index of 1.6 ml/kg
is somewhat greater than that of man.
about 1 ml/kg. However, since the
heart rate was slower in Gigi, the
cardiac index was closer to that of
man (Table 1). Again, in comparing
several species, we note that when
cardiac output is plotted against body
weight on a log-log scale, a straight
line is obtained (Altman and Dittmer,
1971, p. 320). It does seem reason-
able that stroke inde.x is roughly
equivalent in different mammals, since
the heart/body mass ratios are similar.
Although the measurement of
acceleration, as opposed to displace-
ment or velocity, minimizes the in-
fluence of ventilation, any movement
or muscular activity can disturb the
recording. Since whales must expire
and inspire rapidly between dives,
the muscular activity is relatively
violent. As Wahrenbrock has
measured. Gigi's peak instantaneous
flow rate was 285 1/sec. Thus
ventilation demolished Gigi's Beg
recording. Fortunately, ventilatory
rate was extremely slow so that the
Beg had sufficient time to recover
between breaths.
The Beg has now been recorded
in a wider range of masses in animals
than any other physiologic test. The
mass ratio is 1;6, 000.000. egg embryo:
whale. This points out the versatility
of the Beg and suggests its importance
as a technique for comparative
physiological and pharmacological
studies.
ACKNOWLEDGMENTS
The authors wish to acknowledge
the invaluable assistance of Gary
Maruschak. They also thank the
staff of Sea World who participated
in this study. Without their enthusi-
astic cooperation, these studies would
have been impossible.
LITERATURE CITED
AUman. P. L., .ind D. S. Dittmer (editors).
1971. Respiration and circulation. Fed.
Am. Soc. E,\p. Biol.. Bcthescda. 4tO p.
Burger. H. C. and A. Noordergraaf.
19?6. Physical basis of ballistocardiography.
IF The quanties that can be measured
with ditferent types of ballistocardiographs
and their mutual relations. Am. Heart
J. 5 1:127-139.
Burger. H. C. A. Noordergraaf. and A. M.
W. Verhagen. 1953. Physical basis of
the low-fretiuencv ballistocardiograph.
Am. Heart J. 46:71-83,
Cullen. B. F.. E. F Eger. II. N. T. Smith,
D. C. Sawyer. G. A. Gregory, and T. A.
Joas. 1970. Cardiovascular effects of
fluroxene in man. Anesthesiology 32:218-
230.
Deuchar. D. C. 1966. The relations of
hallistixardiography to other methods of
studying the cardiovascular system in
man. In A. A. Knoop (editor). Ballisto-
cardiography and cardiovascular dvnamics,
p. 225-236,' William & Wilki'ns Co..
Baltimore.
Elsb,ich. H., F. A. Rodrigo. H. W. H. Weeda.
and J. Pool. 1970. The ballistocardiogram
in selection for and assessment of physical
conditioning in patients with ischemic
heart disease. Bibl. Cardiol. 26:49-51.
Green. R. F. 1971. Observations on the
anatomy of st>me cetaceans and pinnipeds.
In S. H. Ridgwav (editor). Mammals of
the sea. Biology and medicine. C. C.
Thomas. Springfield, p. 269.
Holloszv. J. O.. J. S. Skinner. A. J. Barry,
and T. K.. Cureton. 1964. Effect of
physical conditioning on cardiovascular
function. Am. J. Cardiol. 14:761-770.
Juznic. G. 1970. The ultra-low frequency
ballistocardiogram of the mouse. Bibl.
Cardiol. 26:280-291.
Leighton. R. F.. A. M. Weissler. P. B. Wein-
stein, and C. F. Wooley. 1971. Right
and left ventricular systolic time inter-
vals. Am. J.Cardiol. 27:66-72.
Moss, A. J. 1961. Ballistocardiographic
evaluation of the cardiovascular aging
process. Circulation 23:434-451.
Noble. M. I. M.. I. T. Gabe. and D. Tren-
chard. 1967. Blood pressure and flow
in the ascending aorta of conscious dogs.
Cardiovasc. Res. 1:9-20.
Noble. M. I. M., D. Trenchard. and A. Guz.
1966. Studies of the maximum accelera-
lit^n of blotid in the ascending .Kirta oi
conscious dogs. In A. A. K.noop (editor).
Ballistocardiography and cardiovascular
dynamics, p. 243-247. William & Wilkins
Co.. Baltimore.
Rushmer. R. F. 1964. Initial ventricular
impulse. A potential key to cardiac
evaluation. Circulation 29:268-283.
1970. Cardiovascular dy-
namics. 3rd ed. W. B. Saunders Co..
Philadelphia, p. 110.
Scarborough. W. R., E. F. Folk. III. P. M.
Smith, and J. H. Condon. 1958. The
nature of records from ultra-low frequency
ballistocardiographic systems and their
relation to circulatorv events. Am. J.
Cardiol. 2:613-641.
Smith. N. T. In press. Ballistocardiography.
In A. M. Weissler (editor). Noninvasive
methods in cardiac evaluation. Grune
and Stratton. New York.
Smith. N. T.. R. L. Van Citters, and P. D.
Verdouw. 1970. The relation between
the ultra-low frequency ballistocardio-
gram, the acceleration pneumocardio-
gram. and ascending aortic flow accelera-
tion in the baboon. Bibl. Cardiol.
26:198-205.
Starr. I., and A. Noordergraaf. 1967.
Ballistocardiography in cardiovascular
research. J. B. llippincott Co.. Phila.,
438 p.
Talbot. S. A., and W. K. Harrison. Jr. 1955.
Dynamic comparison of current ballisto-
cardiographic methods. Part I: Artefacts
in the dynamically simple ballistocardio-
graphic methods. Circulation 12:577-587.
Tannenbaum, O., H. Vessell. and J. A.
Schack. 1956. Relationship of the
natural body damping and body frequency
to the ballistocardiogram. Circulation
13:404-409.
Tischenko. M. 1. 1963. The significance
of natural vibrations of the human body
in shaping the ballistocardiogram. Bio-
physics 8:311-319.
Walker. E. P. 1968. Mammals of the
world. Vol. II. Johns Hopkins Press.
Baltimore, p. 921.
Weissbach. G. H.-J. 1960a. Die Rolle
der Tischmasse und der Wert einer
zusatzlichen Diimpfung beim Elongations-
ballistokardiographcn. [In Ger.) Pflue-
gers Arch. ges. Physiol. 270:529-535.
1960b. Die registrier-
technischen Eigenschaften des direklen
ballistokardiographen. Z. Kreislaufforsch.
49:626-630.
Weissler. A. M.. R. G. Peeler, W. H. Roehll,
Jr., and N. C. Durham. 1960. Relation-
ships between left ventricular ejection
time, stroke volume, and heart rale in
normal individuals and patients with
cardiovascular disease. Am. Heart J.
62:367-378.
Winter, P. J., D. C. Deuchar, M. N. Noble,
and A. Guz. 1966. The ballistocardiogram
and left ventricular ejection in the dog.
In A. A. Knoop (editor). Ballistocardiog-
raphv and cardiovascular dynamics, p.
248-254. Williams c<; Wilkins Co.. Balti-
more.
Winter, P. J., D. C. Deuchar. N. 1. M. Noble,
et al. 1967. Relationship between the
ballistocardiogram and the movement
of blood from the left ventricle in the
dog. Cardiovasc. Res. 1:194-200.
MFR Paper 1046. From Marine Fisheries Review, Vol.
36. No. 4. April 1974. Copies of this paper, in limited
numbers, are available from D83. Technical Informa-
Division. Environmental Science Information
tion
Center. NOAA. Washington. DC 20235.
14
MFR PAPER 1047
Investigation of Blubber Thickness in a
Gray Whale Using Ultrasonography
MICHAEL P. CURRAN and WILLIAM M. ASHER
ABSTRACT
A captive juvenile i;ray whale. Eschrichtius robustus. was studied with tdtra-
soiind using A-mode technique. Measurements of hiuhher and fat titickness by
means of selected tissue interfaces were made. Suture implantation depths were
also measured. Ultrasound woidd he a reliable method for measiirini; hiuhher
and fat thicknesses to .y/ic insiiiht to a marine moiitmal's nutritional status.
PROBLEM
A captive yearling gray whale was
considered for ultrasound study 1) to
measure blubber and fat thickness to
reflect on nutritional status, and 2)
to measure depth of polyethylene
suture implantations being used for an
attachment of a radio transmitter de-
vice on the animal's dorsal surface.
PROPOSAL
Using an ultrasound beam with
A-mode technique, it was proposed to
measure skin, blubber, fat. and muscle
depth. Tissues of varying density will
reflect ultrasound echoes from their
respective interfaces. A porpoise. Tur-
siops truncutus. model was proposed
for correlation.
BACKGROUND
Ultrasound is a relatively new sci-
ence which is meeting with intense
interest and enthusiasm for medical
diagnostic and research purposes. It
has proven effective in detecting brain
midline shifts with the echoencephalo-
gram. Examinations of the heart to
predict cardiac output, mitral valve
activity, and presence or absence of
pericardial effusions are made. B-scan
examination of the abdomen to localize
and characterize various masses and
organs in the peritoneal cavity and
retroperitoneal space is accepted prac-
tice. Obstetrics has found valuable
use for ultrasound in evaluating
gestational age. placental location,
and pelvic masses.
In the field of veterinary medicine
this technique has made it possible
to select breeding stock by determina-
tion of the fat and muscle interfaces,
allowing identification of those ani-
mals with the best commercial poten-
tial. This latter application suggested
measurements for marine animals to
evaluate nutrition.
MATERIALS AND METHODS
Utilizing commercially available
pulsed ultrasound equipment designed
for medical application, multiple
measurements of the echo interfaces
of the gray whale were obtained at
selected positions along the dorsal-
lateral aspect and axilla. Additional
measurements were obtained over the
polyethylene sutures to determine the
Lt. Conidr. Michael P. Curran,
MC. USNR, and Lt. Comdr.
William M. Asher, MC, USN,
are both from the Department
of Ultrasound and the Clinical
Investigation Center, Naval Hos-
pital, San Diego, CA 92134.
The opinions or assertions con-
tained herein are those of the
authors and are not to be con-
strued as official nor as reflecting
the views of the Navy Depart-
ment.
suture depth. All of this material was
displayed on a cathode ray oscillo-
scope with a linear scale divided into
millimeter increments. As an in vitro
correlation to provide information as
to which structures were providing
the echo interfaces observed in the
live mammal, a porpoise model with
necropsy section was obtained. Using
a direct visual placement of the trans-
ducer in similar areas to that of the
gray whale, the echo interfaces were
photographed on the oscilloscope.
Direct linear measurements and ana-
tomical identification of the structures
traversed were performed. These echo
patterns correlated highly with the
similar patterns obtained from the
gray whale and indicated which struc-
tures were providing these echoes.
Thin section radiographs were ob-
tained of the porpoise model, further
demonstrating the density differences
of tissue between the skin lines, blub-
ber, areolar fat, muscle, and fascial
surfaces. In all cases the measurements
corresponded exactly to the visual
interpretation of the fascial, fat, bone,
and skin interfaces.
DISCUSSION
Elementary Ultrasound Physics
Although ultrasonic technology in
medicine is relatively new, the earliest
experiments date back to the I800"s
when attempts to produce high fre-
quency sounds were performed. In
1883 Gallon developed an ultrasound
whistle which was capable of produc-
ing vibrations as high as 25,000 cycles
per second. In modern terminology,
the frequency of vibrations is assigned
the term "Hertz"' and 25,000 cycles
per second is abbreviated as 25 kilo-
heriz (25 kHz). In 1929 Sokolov de-
scribed an ultrasonic method for de-
tecting flaws in metals. Following
this, in l')47. this new modality was
utilized in medical diagnosis when
early workers such as Keksell. in
Sweden, demonstrated the ability to
delect the midline of intracerebral
structures.
15
Electrical to Mechanical
Mechanical to Electrica
Figure 1. — Piezoelectric eftect
Figure 2. — Split function concept. Pulsed ultra-
sound from transducer is transmitted 10% of the
time and received 90% at 400 pulses/sec.
Ultrasonics, the technology of high
frequency sound waves, deals with the
transmission of sound or pressure
waves through a medium. Sound
waves, unlike electromagnetic waves,
cannot be transmitted through a vac-
uum. The generation of sound waves
from a transducer depends on a phe-
nomenon known as piezoelectric ef-
fect. This effect is produced when
electrical energy is applied to a crystal
which, when distorted by this electri-
cal energy, will produce a mechanical
pressure wave. In reverse, the piezo-
electric effect occurs when mechani-
cal energy distorts the crystal, pro-
ducing an electrical potential which
can be measured. The technique of
recording reflected ultrasound results
from this reversible behavior. (See
Figures 1 and 2.) Sound waves travel
through various materials with char-
acteristic velocities. The product of
the density of the material and the
velocity of sound through the given
material is called "characteristic acous-
tic impedance" (Zl. When the two
substances adjacent to each other
transmit sound at a different velocity,
the ultrasonic reflection (/?) at the
boundary is determined by the ratio
of the two acoustic impedances as
described in the formula
A f
^=-f-
1.5 OO
5,000,000 cyC/
If the two substances have the same
acoustic impedance, the numerator
becomes zero and there is no reflection.
On the other hand, if there is a large
difference between the acoustic im-
pedances, the result approaches unity,
and almost all of the energy is reflect-
ed. In between these two extremes
some of the sound energy is reflected
while that remaining passes through
the interface. Since most soft tissues
have acoustic impedances that are
quite similar, there are relatively weak
reflections at the boundaries. The air-
tissue interface is the strongest biologi-
cal reflector. The bone-tissue inter-
face, likewise, produces a very strong
reflection. As most reflections are rela-
tively weak, sensitive equipment is re-
quired to detect those boundaries
with the less strongly reflected echoes
and interfaces such as fat-muscle.
What is the resolution of the sys-
tem? The frequency of sound deter-
mines its wavelength. The resolution
is likewise dependent on the wave-
length in the axial direction. The high-
er the frequenc) . the smaller the
wavelength; thus, we have a better
resolution capabilits as determined
by the formula r = X/ where r is the
velocity of the sound in the medium.
X is the wavelength, and / is the fre-
quency. We assume that the minimum
distance between two objects for dis-
= 3x10"^
= 0.3 mm = A
A X 1.5 = Minimum Distance
Between T\a/o Objects
for Discrimination.
= 0.45 mm For 5 mhz
Figure 3. — Formula lor resolution.
crimination must be equal to at least
l'/2 wavelengths. (See Figure 3.) By
going to higher frequencies, however.
we lose penetration in tissue due to
sound attenuation: therefore, a com-
promise must be made and the fre-
quency selected which gives adequate
axial resolution, yet adequate penetra-
tion through the tissue thickness. For
the mammalian models studied the
frequency varied between 1 and 2
megahertz, which was adequate for
penetration through the structure
studied.
Findings
Although the size dilterence be-
tween the 28 foot captive gray whale
and the captive Atlantic bottlenosed
dolphin {.Titrsiops nuncaliis) necrop-
sy model is somewhat different, the
anatomical structures of the mammals
are known to be similar.
During periods of illness or mal-
nutrition, marine mammals of these
species are noted to develop a depres-
sion in the dorsal contour posterior to
the axilla. It is thought that this de-
pression is due to catabolism of blub-
ber, areolar fat. and/or muscle mass
loss.
Presuming ihat areolar fat dimin-
ishes in volume prior to muscle loss.
Figure 4. — Radiograph of Turs/ops cross sec-
tion demonstrating (1) blubber. (2) areolar lat.
(3) muscle group to fascial layer, and (4) second
deep muscle group to dorsal spinous process.
16
17
18
one could measure the normal thick-
ness in heahhy mammals and compare
with abnormal animals and any avail-
able necropsy specimens.
As a pilot program, echographic
measurements were made in both the
gray whale. E. robitstus. and the
porpoise. T. nunccUiis. Necropsy cor-
relation in Tursiops showed that the
measurements were easy to perform
and were highly accurate. (Figures
4-8.)
Applications
Figure 6. — A-scan of Tursiops
cross section. Lettered spikes
conform to radiograpfis and
tissue boundaries as measured
in necroscopy section: (A-
B) blubber ttiickness = 2
cm. (C-D) fat thickness =
1 cm, and (D-E) muscle
ttiickness = 8.2 cm.
Ideally, to make this method most
useful, measurements should be made
and necropsy correlation measure-
ments obtained whenever these mam-
mals are found deceased. Due to the
expense and shortage of the species,
flying a small team to the animal site
with the easily portable battery or
generator operated scanning equip-
ment should be the most effective
means of collecting this invaluable
data.
Further observations on the nutri-
tional status during development com-
paring captive and free animals, as
well as disease effects, should prove
to be a new approach to the study of
marine mammals. Such research data,
if accumulated, may be of great bene-
fit in the protection and treatment of
valuable, trained marine mammals
and their free swimming counterparts.
CONCLUSION
A-mode echography is an effective
means of measurement of tissue layers
and should be an effective tool in the
study of marine mammal nutrition
and health status.
Figure 7. — A-scan of gray
wtiale dorsal-lateral surface
posterior to axilla demonstrat-
ing blubber thickness of 4.1
cm (A-B) and fat thickness of
4.6 cm B-C). C represents
fat-muscle interface.
Figure 8. — A-scan of gray
whale for polyethylene suture
localization. (A) Skin. (B)
Blubber-fat interface at 3.5
cm. (C) Polyethylene suture
at 5.5 cm. (D) Fat-muscle
interface at 7 cm.
Figure 5. — Radiograph of Tursiops cross section
demonstrating (A) skin-blubber interface, (B)
blubber-fat interface, (C) fat-muscle interface.
(D) muscle-fascial layer interface, and (E) re-
flective bone (dorsal spinous process).
19
REFERENCES
Ashcr, W. M.. und A. K.. Freimanis. 1%^.
Echographic diagnosis of retroperitoneal
lymph node enlargement. Ultrasound scan-
ning technique and diagnostic findings.
Am. J. Roentgenol. Radium Ther. Nucl.
Med. 105:438-445.
Freimanis. A. K., and W. M. Asher. 1970.
Development of diagnostic criteria in
echographic study of abdominal lesions.
Am. J. Roentgenol. Radium Ther. Nucl.
Med. 108:747-755.
Grossman, C. C, J. E. Holmes. C. Jovner,
and E. W. Purnell (editors). 1966. Di-
agnostic ultrasound. Proc. First Int. Conf.
Univ. Pittsburgh, 1965. Plenum Press,
New York, 5 19 p.
Lehman, J. S. 1969. Ultrasonography.
Delaware Med. J. 40:24-25.
Okasawa, A., and L. Hakkinen. 1969.
Comparative experiments on the attenua-
tion of ultrasound in muscular and fat
tissue. Acta Aphthal (Kohenhavn) 47:735.
Strakova, M.. and J. Markova. 1971. Ul-
trasound use for measuring subcutaneous
fat. Rev. Czech. Med. 17:66-73.
MFR Paper 1047. From Marine Fisheries Review, Vol.
36, No. 4, April 1974. Copies of this paper, in limited
numbers, are available from D83, Technical Information
Division, Environmental Science Information Center,
NCAA. Washington. DC 20235.
MFR PAPER 1048
MATERIALS AND METHODS
Surgical Attachment of a Telemetry Device
to the Dorsal Ridge of a Yearling
California Gray Whale, Eschrichtius robustus
JOHN C. SWEENEY and JOEL L. MATTSSON
ABSTRACT
Siiri^iccil atuichiiu'iil of an instrument packufie moiiniinf^ device onto the dorsul
rUli;e of a yenrliiii' feinnle California i-ray whale. Eschrichtius robustus, wa.\
accomplished lhroui;h the utilization of four lariie polypropylene sutures. Use
of polypropylene and polyester fabric meshes to induce tissue i;ronth aroutid the
sutures was not successful. Post-operative therapy was heneficial in iiisuriui.;
adequate healing at the suture sites. The original polypropylene sutures were
replaced the day before release by polyvinyl chloride coated stainless steel.
INTRODUCTION
In March 1971, an infant female
gray whale was captured within
Scammon's Lagoon, Baja California,
and subsequently transported by boat
to Sea World, Inc. in San Diego. Calif.
The animal was captured for research
purposes, and for the year following
her capture, various studies were un-
dertaken.
As the animal approached 1 year of
age. the financial burden to Sea World
in holding facilities, personnel, and
food made it necessary to design a
plan for her release. At that time.
W. E. Evans, of the Naval Undersea
Center. San Diego, proposed (with the
support of the National Oceanic and
Atmospheric Administration) that the
whale be released carrying a telemetry
device for tracking and recording.
Evans (1971) has reported the use
of radiotelemetry devices attached to
the dorsal fin of dolphin, using a bolt
placed through the tin. Martin. Evans,
and Bowers (1971) have utilized a
harness for the fixation of a device
onto a pilot whale. A gray whale has
no dorsal fin for bolt fi,\ations. and
the growth rate of this animal left the
harness method undesirable. There-
fore, a surgical fixation was considered
the method of choice.
J«»hn C. Sweeney and Joel L.
Maltsson are associated with the
Naval Undersea Center, San
Diego, C A 91132.
Sutures composed of .■* mm diame-
ter polypropylene were swaged onto
a stainless steel needle made from 3
mm diameter rod shaped into a 10 cm
diameter half circle. Pohpropylene
was chosen because of its inert nature
in mammalian tissues (Usher et al.,
1962) and because of its availability
in the dimensions required. Two types
of prosthetic mesh were used in con-
junction with the sutures, polypro-
pylene (Marlex®') mesh and polyester
fiber (Mersilene*-).
Five weeks before the scheduled re-
lease, an attempt was made to place
polypropylene mesh pads (2 cm X 2
cm) subdermally at the entrance and
exit sites of the four proposed sutures
at positions on a longitudinal plane
10 cm to either side of the dorsal
ridge and 10 cm apart. The intention
was to induce collagen fiber infiltra-
tion within the fabric to add strength
to the skin and to prevent infiltration
of water once the sutures were in place.
The skin was closed with simple inter-
rupted nylon sutures.
Four weeks before release the four
polypropylene sutures, each having
had a sheet of polyester fabric at-
tached to it using Eastman 9-10 ad-
hesive.'' were placed at the proposed
sites. Depth of penetration of the su-
tures was later confirmed by ultrason-
ography to be from 4 to 6 cm (Curran
'Cavol. Inc., Providence. R.I. Reference to trade
names does not imply endorsement by the Na-
tional Marine Fisheries Service, NCAA
-Ethicon. Inc , Scmerville, N J
^'Eastman Chemical Products, Kingsport, Tenn,
20
and Asher. 1974) all King within the
I'attN tissue between blubber and
muscle. Once in position, the suture
ends were temporarily fused, each
suture forming a ring enclosing the
dorsal ridge (Figure 1). Each surgical
procedure was done under local an-
esthesia, using 2 percent Xylocaine.
RESULTS
Both attempts to utilize mesh fabrics
were unsuccessful. Because no fascial
interface is present between epidermis
and dermis, or dermis and hypodermis.
in cetaceans, placement of mesh pads
under the skin was not accomplished.
After attempts were made at two of
the operative sites, it became apparent
that it would be too difficult to embed
the pads properly. In addition, the
sutures cut through the epidermis
when even light tension was applied,
preventing adequate closure of the
incision. Because of these problems,
the procedure was not completed.
Within 5 days, each of the mesh pads
had been sloughed.
The mesh coated sutures did not
induce tissue infiltration, but rather,
acted as an irritant with a consequent
tissue inflammatory response.
Some drainage from the suture
holes was observed on the third post-
operative day. and at this time, all
four sutures were easily moved back
and forth within their tissue bed. The
exudate was composed of clear, non-
viscous fluid containing tags of white
coagulated matter dispersed through-
out. Cellular composition was 70 per-
cent mature neutrophils and 30 per-
cent lytnphocytes. Swabs were taken
on the third postoperative day and on
two subsequent occasions. No bacteria
were found. Daily flushing of each
suture site with normal saline and
nitrofurazone solution was done for
the ne\t three postoperative weeks. At
no time did the animal appear sick,
nor was there any indication in her
blood tests to suggest that an infection
Figure 2. — Normal healing around polypropylene
sutures.
Figure 1. — Polypropylene sutures in position with ends (used, forming a ring
enclosing the dorsal ridge.
21
was present. By the end of the 3 week
postoperative period, normal healing
was considered well underway (Figure
2). and there was. by then, no drainage
from any of the suture sites, though
the sutures were still freely movable.
One week before the scheduled re-
lease, the instrument package saddle
was mounted onto the sutures to allow
the animal time to adjust to it before
adding the somewhat heavier (approxi-
mately 6 kg) instrument package itself.
The animal occasionally rubbed the
saddle against the side of the tank
until the attachment was tightened to
reduce free-play of the saddle as the
animal swam. On the day before re-
lease, cracking of the polypropylene
sutures was noticed, requiring their
replacement with sutures of the same
diameter composed of polyvinyl chlor-
ide coated stainless steel. These were
found to be more pliable and stronger
than the polypropylene.
At the last visual sighting of the
animal on 7 April 1972. the instru-
ment package was still securely at-
tached despite the fact that, on several
occasions, kelp had been seen trailing
from it (J. S. Leatherwood. pers.
comm.). At this time, we have no in-
dication that this procedure has. in
any way. compromised the ability of
this animal to survive.
LITERATURE CITED
Curran. M. P., and W. M. Asher, 1974.
Investigation of blubber thickness in a
gray whale using ultrasonography. Mar.
Fish. Rev. 36(4): lS-20.
Evans. W. E. 1971. Orientation behavior
of delphinids: radio telernetric studies.
Ann-N.-*'. .Acad. Sci. 188:142-160.
Martin, H., W. E. Evans, and C. A. Bowers.
1971. Methods for Radio Tracking Ma-
rine Mammals in the open sea. IEEE Eng.
in the Ocean Environ. Conf. p. 44-49.
Usher, F. C, J. E. Allen, Jr., R. W. Crosth-
wait, and J. E. Cogan. 1962. Polypro-
pylene monofilament. A new biologically
inert suture tor closing contaminated
wounds. Am. Med. Assoc. 179:780-782.
MFR Paper 1048. From Marine Fisheries Review, Vol.
36, No. 4. April 1974. Copies of this paper, in limited
numbers, are available from D83. Technical Informa-
tion Division, Environmental Science Information
Center, NCAA, Washington, DC 20235.
MFR PAPER 1049
Some Hematologic Observations on
the California Gray Whale
ALFRED ZETTNER
ABSTRACT
E.Kiiininalitin of llic blood of the California i;ray whale, ohlaiiwil .shortly
after its arrival at Sea World, San Dieijo revealed the followiiti; data: H'BC-IJI.^
X \0->lciihic mm: RBC-2.4 X IO<^jcuhic mm: hemof^lohin-IO.O gjlOO ml:
hematocrit-31 percent: MCV-128 jj-^: MCH-42.8 tijdii: MCHC-32.4 percent.
Hemonlohin electrophoresis showed a sini;le hemof^lohin hand with a mobility
similar to that oj human hemoi;lobiii /•'. I he whale hemoiilobin was 100 percent
alkali resistant. No changes of this hemoi;lobin were seen on repeated analyses
over the course of 12 months.
The capture of a young, female
California gray whale. Eschrichiiits
rohitstiis, in .Scammon's Lagoon, and
its maintenance in captivity at Sea
World, San Diego for 12 months pro-
Alfred Zetlner is a physiciun
with the Division of Clinical
Palholouy, Depart incnt of Palii-
olosy. School of Medicine, L'ni-
versitv of California, San Diego,
CA 92103.
vided the opportunity for some hema-
tologic studies which are to be report-
ed here.
ROUTINE BLOOD
EXAMINATION
A heparini^ed blood sample ob-
tained on 18 March 1971. one day
after the arrival of the whale at Sea
World, was brouuht to the Clinical
Laboratories of University Hospital.
University of California. San Diego.
The blood was analyzed on a Coulter
Counter.' Model "S". which allows
the automatic simultaneous determin-
ation of cell counts, mean corpuscular
volume (MCV). and hemoglobin con-
tent. The hematocrit, mean corpuscu-
lar hemoglobin (MCH). and the mean
corpuscular hemoglobin concentra-
tion (MCHC) are automatically com-
puted from the three parameters
measured (Pinkerton et al., 1970)
The instrument is standardized twice
daily and performs approximately 200
analyses per day for clinical purposes.
The results were the following:
WBC-13.9 X lO^Vcubic mm
RBC-2.4 X lO'Vcubic mm
Hemoglobin- 1 0.0 g/ 1 00 ml
HCT-3 I percent
MCV- 128 /J-'
MCH-42.8;ttiUg
MCHC-32.4 percent
A blood smear was prepared and
stained by the automatic HEMA-
TEK- technique, which employs a
'Coulter Electronics, Inc , Hialeati. Fla Refer-
ences to trade names does not imply endorse-
ment by ttie National tvlarine Fistienes Service.
NOAA.
^Ames Company. Division of Miles Laboratories,
Inc. Elktiart. Ind
22
moditicd Wrighl-Giemsa slain, and
examined by oil immersion microscopy.
The red cells were round, moderate-
ly anisocytotic ranging from 7.5-9.5 /i
in diameter, and appeared well hemo-
globinated with only occasional slight
central pallor. An occasional red cell
displayed polychromasia. and some
rare Howell-Jolly bodies were seen.
No nucleated red cells were encoun-
tered.
A white cell differential count was
as follows;
Segmented neutrophils b^ percent
Band forms 19 percent
Metamyelocytes < 1 percent
Monocytes 9 percent
Lymphocytes 8 percent
No eosinophils nor basophils were en-
countered. The lymphocytes were all
of the large type. No small lympho-
cytes with the typically scant cyto-
plasm and dark staining nuclei were
present. Twenty-one percent of the
nuclei of the mature, segmented neu-
trophils had distinct '"drumstick" ap-
pendages.
The thrombocytes appeared as
round platelets, with diameters ap-
proximately one-third to one-half of
those of the red cells. Their number,
estimated from their frequency distri-
bution on the smear in relation to the
erythrocytes, was in the range of
300.000-350,000/cubic mm.
HEMOGLOBIN
ELECTROPHORESIS
Hemoglobin electrophoresis was
performed by the vertical acrylamide
gel technique as described in detail
elsewhere (Bierman and Zettner. 1967)
(Nakaniichi and Raymond. 1963).
Briefly, a toluene hemolysate of the
washed red cells is prepared and elec-
trophoresed in Tris-buffer of pH 9,0
for 3'/2 hours at 120 ma. The acryla-
mide gel slabs are then stained with
amido black and destained electro-
phoretically in 5 percent acetic acid.
The results are shown in Figure 1.
The whale hemoglobin (Slots Nos. I
and 7) migrated slightly slower than
human hemoglobin A. The position
SLOT I
3 4 5 6 7
6-9'71
Whale non-heme protein
(Whale Carbonic Anhydrase ?)
Human Corbonic Anhydrase B
Humon HbAg
Human HbS
Gray Whale Hb
Human HbA
fill •
Figure 1. — Vertical acrylamide gel hemoglobin electrophoresis. Tris-buffer, pH 9.0. The gel contains
the toluene hemolysates of the following: Slots No. 1-gray whale; No. 2-human with A-S trait; No. 3, 4-
normal humans; No. 5.6-standards; No. 7-gray whale (same as slot No. 1). Original (sample
application slots) at top. Cathode-lop; anode-bottom.
of the band of the whale hemoglobin
was indistinguishable from that where
human hemoglobin F-' would be ex-
pected. No minor hemoglobin com-
ponents equivalent to those found in
human blood could be detected. The
weakly stained band of much slower
mobility, as shown in Figure I, is a
non-heme protein, as indicated by the
failure of this protein band to react
with benzidine when a freshly electro-
phoresed, unstained strip of the gel
containing the whale sample was sub-
mersed in a benzidine and peroxide
solution.
The pattern of hemoglobin electro-
phoresis performed on blood samples
obtained on 17 March and 27 April
1971, and 13 March 1972 was iden-
tical to that demonstrated here.
ALKALI DENATURATION
A quantitative alkali denaturation
test performed on the toluene hemoly-
sate by the method of Singer, Chernoff,
and Singer ( 195 1) revealed the whale's
hemoglobin to be 100 percent alkali
resistant. The alkali resistance of the
hemoglobin was the same in all sam-
ples obtained over the course of 1
year, as listed above.
DISCUSSION
The values of the various red cell
parameters, as reported here, are in
fair agreement with those published by
Lenfant (1969). Relative to most ter-
restrial mammals, the California gray
whale appears to have lower red cell
counts, hemoglobin concentrations,
and hematocrits, although the MCV
is considerably in excess of 100 jU^.
A proportional increase of the MCV
of red cells with total body length of
marine mammals of different species
has been shown (Lenfant. 1969). Of
interest is the finding of Lenfant ( 1969)
of a high proportion of nucleated red
cells in the gray whale. This is in
distinct contrast to the complete ab-
sence of nucleated red cells in the
blood samples examined here. It
should be considered that the previous
observations were apparently made on
sick, wounded, dying, or dead animals:
and that under these abnormal condi-
tions, immature red cells may have
been released into the circulation.
The only indication of young red cells
in our samples were the rare Howell-
Jolly bodies and occasional polychro-
masia.
23
The white cells were remarkable in
that no small lymphocytes, eosinophils,
or basophils were seen. Otherwise,
their numbers and percentages appear
to be near the normal limits. Of interest
is the occurrence of "drumstick" ap-
pendages in 21 percent of the mature
segmented neutrophils. These were
described by Davidson and Smith
(1954) in human blood as a genetic
sex indicator for females. They occur
in 1-17 percent of the segmented
neutrophils of all human females and
are thought to represent the inactivat-
ed X-chromosome. analogous to the
Barr body observed in most somatic
cells. It can be reasonably assumed
that also in the whale, drumsticks in
the neutrophils are indicators for the
female sex.
The uniform electrophoretic mobil-
ity of this gray whale's hemoglobin,
characteristic of human hemoglobin
F. is in accordance with the finding of
others (Lenfant, 1969). Of further
interest was the hemoglobin's resistance
to alkali denaturation. However, no
conclusions can be drawn from this
coincidental sharing of two physical
properties with human hemoglobin F
as to functional or structural similari-
ties between these two hemoglobins.
The reasons for the alkali resistance
of certain hemoglobin variants are
poorly understood. In the human this
is related not only to the presence of
gamma chains in the hemoglobin
molecule, but also to the structural
relationships of the various chains to
each other. For instance. Bart's hemo-
globin, composed of four gamma
chains, is only half as alkali resistant
as hemoglobin F, which is a tetramer
of two alpha and two gamma chains.
The elucidation of the structure of
the gray whale's hemoglobin depends
on the full analysis of its amino
acid sequence. .Such an undertaking
can also be expected to provide some
evolutionary clues for the California
gray whale.
From the evidence presented here,
it appears that this species possesses
only one type of structurally uniform
hemoglobin, although the possibility
that we are dealing with two or more
hemoglobins of identical electropho-
retic mobility and alkali resistance
cannot be entirely excluded.
The band of non-heme protein ap-
pears to be analogous to a similar
band which is consistently seen in
the electrophoretograms of human
bloods. In the latter, this is known to
represent carbonic anhydrase B. a red
cell constituent persistently extracted
with the toluene hemolysates.
LITERATURE CITED
Bicrman, A. H.. and A. Zettner. 1967. A
simple electrophoretic method for the
quantitative determination of hemoglobin
A2. Am. J. e'lm. Palhol. 48: 1.19-146.
Davidson. W. M.. and D. R. Smith. 19.S4.
A morphological sex difference in the
polymorphonuclear neutrophil Icukocvtes.
Brit. Med. J. ::6-7.
Lenfanl, C 1969. Physiological properties
of hlood of marine mammals. In H. T.
Andersen (editor). The biology of marine
mammals, p. 95-1 16. Academic Press. N.Y.
Nakamichi. M.. and S. Raymond. 196,1.
Acrvlamide-gel electrophoresis of hemo-
globins. Clin. Chem. 9: U.S. 145.
Pmkerton, P. H., I. Spence, J. C. Ogilvie,
W. A. Ronald. P. Marchant, and P. K.
Ray. 1970. An assessment of the Coul-
ter counter model S. J. Clin. Palhol. 23:
68-76.
Singer. K., A. I. Chernoff. and L. Singer.
1951. Studies on abnormal hemoglobins.
1. Their demonstration in sickle cell anemia
and other hematological disorders by
means of alkali denaturation. Blood J.
Hematol. 6:413-428.
MFR Paper 1049. From Marine Fisheries Review, Vol.
36, No. 4. April 1974. Copies of this paper, in limited
numbers, are available from D83, Technical Information
Division. Environmental Science Information Center,
NOAA. Washington, DC 20235.
MFR PAPER 1050
Some Coagulation Factors in Plasma from a
California Gray Whale, Eschrichtius robustus
W. MEDWAY
ABSTRACT
.4 cilniled pkisimi sample was assayed for some couiiuhnion faelors. The
levels ohiaineil were compared with those from some of ihe small tooilied
whales. Factor XII activity was verx low in the i^rav whale sample, whereas
toothed whales have none.
INTRODUCTION
Many people working with small
odontocete whales in captivity have
made the observation that whale
blood has a prolonged clotting time.
Since this observation was made two
reports have described the lack of
clotting Factor XII in blood in some
of the smaller whales (Lewis, Bayer,
and Szeto, 1969: Robinson, kropat-
kin, and Aggeler, 1969). Another pub-
lication reports a prolonged clotting
time of blood from other small whales:
however, assays for Factor XII were
not made (Ridgway, 1972). There
were no reports of similar studies on
blood from any baleen whale: hence
this report on some studies on a plas-
ma sample from a captive California
gray whale. Eschrichtius rohiistiis.
24
MATERIALS AND METHODS
A titrated plasma sample was ob-
tained from a young ( 1-2 years) female
California gray whale kept in cap-
tivity in San Diego. Calif. The sample
was deep-frozen and shipped via air
express to Philadelphia where the
assays were made. The plasma sample
was slighth llpcmic. The prothrombin
time, partial thromboplastin time.
Factor V. Factor XI, and Factor XII
assays were made in the Coagulation
Laboratory at the Hospital of the
UniversitN of Pennsylvania. It was not
possible to do a fibrinogen assay on
the sample.
Standard laboratory procedures em-
pkning commercial reagents were
used to conduct the assays, with the
e.xception of Factor XII where dol-
phin, Tiirsh'ps iniiucims. plasma was
used as the substrate. Plasma reagent
from Factor XI deficient cattle was
used for the Factor XI assay.
RESULTS AND DISCUSSION
The results of the assays on the
gray whale plasma and some results
on a few odontocete whales, from the
literature, are shown in Tabic I .
The divergence of our results on
the gray whale plasma for prothrom-
bin time, partial thromboplastin time.
Factor V, and Factor XI assays from
those of the two species of odontocete
whales can be explained perhaps on
the elapsed time between sampling
and assay. The presence of a low level
of Factor XII in the gray whale plas-
ma to the non-existence in odontocete
plasma warrants some consideration.
The significance of this difference
teleologically is not known. One of
the problems encountered by deep
diving humans is decompression sick-
ness. This sickness is attributed to the
formation of microdots (disseminated
W. Medway Is associated with
the Department of Clinical
Studies, School of Veterinary
Medicine, University of Pennsyl-
vania, Philadelphia, PA 19104.
Table 1. — A comparison of some clotting factors between odontocete whales and a baleen whale
The numbers in parentheses indicate the number of samples.
Tursiops
Orcinus
orca-
Eschnchtius
robustus
Prothrombin time (sec)
Partial thromboplastin time (ptt) (sec)
Factor V(%)
Factor XI (%)
Factor XI t {%)
17.0(14)
15.6(3)
26.5(1)
346 (15)
216 (3)
107 (1)
136 (14)
239 (3)
17 sec (1)
92.7(14)
146 (3)
24.6(1)
0 (15)
0 (3)
3.4(1)
' Lewis, Bayer, and Szeto (1969)
- Robinson. Kropatkin, and Aggeler (1969).
■1 Ridgway (1972).
intravascular coagulation) with re-
sulting consequences. It is known that
slow-moving acid blood has a pro-
pensity to clot faster. This property
has been attributed to activation of
Factor XII and subsequent clot for-
mation.
Whales dive deeply and are not be-
lieved to suffer from decompression
sickness. Perhaps the lack of Factor
XII or low levels of it is nature's way
of protecting the animals.
ACKNOWLEDGMENTS
The author wishes to express appre-
ciation to H. A. Wurzel. Director.
Coagulation Laboratory. Hospital of
the University of Pennsylvania, and
his staff as well as to J. C. Sweeney,
Naval Undersea Center, San Diego,
Calif, for providing the opportunity to
make this study on the gray whale.
LITERATURE CITED
Lewis. J. H., W. L. Bayer, and I. L. F. Szeto.
1969. Coagulation factor XII deficiency
in the porpoise. Tursiops Inimatits. Comp.
Biocheni. Physiol. .M:667-(-70.
Robinson. A. J., M. Kropatkin. and P. M.
Aggeler. 1969. Hageman factor (factor
.XII) deficiency in marine mammals. Sci-
ence (Wash.. D. C.) 166:14:0-14::.
Ridgway. S. H. 197:. Homeostasis in the
aquatic environment. In S. H. Ridgway
(editor). Mammals of the Sea. p. 65.^.
Charles C. Thomas. Springfield.
MFR Paper 1050. From Marine Fisheries Review. Vol.
36. No. 4. April 1974. Copies of this paper, in limited
numbers, are available from D83, Tecfinical Informa-
tion Division, Environmental Science Information Cen-
ter. NOAA. Wasfiington. DC 20235.
MFR PAPER 1051
Fluorescent Karyotype of the
California Gray Whale
DEBORAH A. DUFFIELD
ABSTRACT
The Jhion-.sccnl kuryolypc of the California .i;ray nhale. Eschrichtius robustus,
is presented and the aae of the fluorescent handini; tecltniqiie for distini;uishini,'
between various cetacean karyotypes is discussed.
The California gray whale. E\ch- (Benirschke's unpublished data cited
ricliiiiis rohitstiis (Gibbosus) has a in Kulu, 1972; Arnason. 1972). Since
diploid chromosome number of 44 reporting of the gray whale karyotype.
25
B.
U
X X
il H M H n
if IS II ti II
II It II tt II It
Figure 1. — Karyotype of the California gray wfiale, Gigi. The autosomes are arranged into four
groups based on centromere positron and relative size. Tfie provisional X ctiromosomes are
indicated.
prepared by standard honiogeneou.s
staining techniques, advances in the
differential staining of chromosomes
have added another dimension to
tcaryotypic analysis by making it now
possible to individually characterize
each chromosome of the complement.
Consequently, and as part of a larger
cytotaxonomic study of marine mam-
mals, evaluation of the gray whale
karyotype by quinacrine mustard
fluorescent banding was undertaken
on Gigi. a captive female gray whale.
MATERIALS AND METHODS
Chromosome preparations were ob-
tained by blood culture (Kulu. Veo-
melt. and Sparkes. 1971). Exposure
of the cells to 0.075 M KCI for 8
minutes was the preferred hypotonic
treatment and cold, rather than tlame-
dried. slides were made. The slides
were stained with Giemsa for normal
karyotyping or with quinacrine mus-
tard (50 micrograms/ml buffer for
.^0-40 minutes) for fluorescent karyo-
typing. Photographs of fluorescent
metaphases were taken on Kitdak'
Tri-X film with an exposure time of
45-50 seconds. Ten Giemsa and eight
fluorescent karyotypes were analyzed.
' Use of trade names m this publrcation does
not imply endorsement ot commercial products
by ttie National Marine Fistieries Servrce
RESULTS
Gigis karyotype (Giemsa) is illus-
trated in Figure 1. The autosomes are
provisionally arranged into four
groups (designated A, B, C. and D).
Group A is composed of five pairs of
large submetacentric chromosomes.
Group B of five pairs of medium-sized
submetacentrics. Group C of six pairs
of metacentrics, and Group D of five
pairs of acrocentric chromosomes.
Within each group the chromosome
pairs are arranged by decreasing size.
The presumptive .\ chromosomes are
indicated in the karyotype.
The fluorescent karyotype of Escli-
ritiniiis rohuslus is presented in Figure
2. The arrangement of the chromo-
somes follows that of the standard
karyotype. The banding pattern of
each chromosome pair is distinctive
and in addition to allowing positive
identification of the homologues
makes it possible to characterize each
pair of the complement in order to
facilitate karyotypic comparison with
other species. The fluorescent banding
pattern of the presumptive X chromo-
Dehorah A. Duftield is with the
Deparlmenl of Biology at (he
University of California at Los
Angeles, Los Angeles, C A 90024.
some is the same as that exhibited by
the X chromosome of another of the
baleen whales, the sei whale. BaUwiiop-
icrii horcalis-, and a number of the
smaller odontocete cetacean species
(personal observation).
DISCUSSION
The karyotype of the California
gray whale appears to be very similar
in number and gross morphology to
that of a number of other cetaceans,
both mysticete and odontocete'' (Kulu.
1972: Arnason. 1972). The fluorescent
karyotype of the gray whale was ex-
amined in the hope that the resolution
of chromosome structure afforded by
fluorescent banding would indicate
differences between its karyotype and
that of other cetaceans not obvious by
regular staining methods. In order to
illustrate the level of karyotypic com-
parison made possible by fluorescent
banding, the larger submetacentrics
which comprise Group A (pairs 1-5)
in two mysticete and two odontocete
species are shown in Figure 3. While
there are certain similarities between
the banding patterns of all four species,
it is clearly possible to distinguish
between the overall banding pattern
of the mysticete (gray whale, sei
whale) chromosomes and that of the
odontocetes ['l'uisi<>p\ inincaius. Laf--
cnorhyiu hits tihliqiiidciis). Less obvi-
ous differences are also present which
further distinguish gray whale from sei
whale and Tiirsiops from Lugeno-
rhyncluis. A detailed comparison of
the fluorescent karyotypes of these
cetaceans is beyond the scope of this
report; however, it can be concluded
- Arnason (1972) tias reported ttiat ttie X ctiromo-
some of B boreahs is one ot ttie larger ctiromo-
somes of ttie complement, sucti as found in
other ot the balaenopteran whales However,
both standard and fluorescent karyotypes ot a
male sei whale, tissue from which was made
available to this author by the Richmond whaling
station in California, indicate that the X chromo-
some of 8. boreahs is of medium size and
similar both in relative size and banding pattern
to the provisional X chromosome of the odon-
tocetes (Kulu, 1972),
■1 Of all cetaceans studied to date, only the
sperm, pigmy sperm, and killer whales are
karyotypically distinct by standard staining
techniques.
26
Figure 2. — The fluorescent karyotype of the California gray whale. Note (hat the members of each
pair have identical bands while one pair can be distinguished from any other by its characteristic
banding pattern.
Figure 3. — Comparison of the banding patterns of four species. Only group A chromosomes are
shown and only one chromosome from each pair per species. Species 1 and 2 are mysticetes,
3 and 4 are odontocetes.
27
from these initial observations that the
comparison of fluorescent banded
karyotypes will significantly enhance
the potential contribution which kary-
otypic analyses can make to the reso-
lution of the phyletic interrelationships
of the modern Cetacea.
ACKNOWLEDGMENTS
My thanks to Sea World and Jay
Sweeney of the Naval Undersea Cen-
ter. San Diego. Calif, for making the
gray whale samples available to me.
The fluorescent technique followed in
the preparation of the karyotypes is
that of Helga Muller. UCLA Medical
Genetics Unit. Los Angeles. Califor-
nia. This work was supported by a
Mental Retardation Training Grant
from the Departments of Psychiatry
and Medicine, UCLA Medical School.
Los Anceles. California.
LITERATURE CITED
Arnason. U. 1972. The role of chromo-
siimal rearrangement in mammalian speci-
alion with special reference lo Celacea
and Pinnipedia. Heredilas 70:113-118.
Kulu. D. D. 1972. Evolution and cyto-
genetics. //; S. H. Ridgway (editor).
Mammals of the sea. Biologv and medi-
cine, p. 503-.'i27. Charles C. Thomas.
SpringHeld.
Kulu, D. D., I. Veomctt. and R. S. Sparkes.
1971. Cytogenetic com.parison of four
species of cetaceans. J. Mammal. 52'
828-832.
MFR Paper 1051. From Marine Fisheries Review, Vol.
36, No. 4, April 1974. Copies of this paper, in limited
numbers, are available from D83. Tectinical Information
Division, Environmental Science Information Center,
NCAA. VJashington. DC 20235.
MFR PAPER 1052
Some Physiological Parameters of the
Blood of the California Gray Whale
WILLIAM G. GILMARTIN, RICHARD W. PIERCE,
and GEORGE A. ANTONELIS, JR.
ABSTRACT
HciihUcK ril. (I Oo-Hh iliwiniiUinn curve, unci hlood vahinic luivc hccn clctcr-
niiiicil fur a Ccilifoniiti .y/'d.v wIhiIc, Eschrichtius robustus. ninl ihc results arc
compcircci lo sonic physioloiiical hlood properties of other celncccins. The
E. robustus hii\ n hlooil volume that is similar lo values esiiiinited for lari;e
whales hy oilier iiiilhors. This is the firsi lime isolopie teehnicjiies have been used
to determine a lariie cetacean's hlood volume.
Large cetaceans do not appear to follow the trend of most lerreslerial mammals
when the body size and P50 ure compared. The P50 /'"' the E. robustus was 36.5 mm
Hi; and is the hii>hest reported for any cetaccdii.
The determination of the physio-
logical properties of the blood of large
cetaceans has been contined primarily
to animals that are stranded or have
been dead for many hours before
blood samples can be drawn. Lenfant
(1964) has summarized most of the
data available on marine mammals.
The capture and maintenance of
Gigi. a California gray whale. E\ch-
richliiis rohiistiis. has given us the
opportunity, for the first time, to study
a large cetacean under definable con-
ditions and to determine its blood
volume and oxygen-hemoglobin dis-
sociation curves.
METHODS
On two separate occasions the
whale was given lO/jCi of radioiodin-
Buth William G. Giliiiartin and
George A. Antunelis, Jr. are
with the Naval Undersea Center
Bio-Svstenis Program, San Di-
ego, "CA 92132. Richard W.
Pierce is with the Coastal Ma-
rine Laboratory, Division of
Natural Science, University of
California. Santa Cruz, CA
95060.
ated human serum albumin (Risa)'.
The labeled compound was adminis-
tered to the animal in one of the
brachial vessels in the right pectoral
tin. Two blood specimens were taken
following each determination to insure
that mixing was complete and the
albumin was not being eliminated
rapidly from the serum. In the first
test (27 December 197 1 1 blood vol-
ume determination samples were taken
at 14 and 20 minute intervals and in
the second test (6 March 1972) were
collected at 10 and IX minute intervals
after administration of (he labeled
compound. The blood samples re-
moved tor counting were taken from
one oi the brachial vessels of the left
pectoral lin and put into well-hepann-
ized tubes. Three ml of the heparinized
whole hlood was added to .^ ml of I
' Abbott Laboratories, Chicago, Illinois. Use
of trade names in this publication does not
imply endorsement of commercial products
by the National fvlanne Fisheries Service.
28
Figure 1. — Oxyhemoglobin dissociation curves
for three cetaceans [T. gilti. G. scammoni, E.
robustus). All curves have been corrected for
pH — 7.4 and determined at 37~C.
percent acetic acid to iyse the red
cells to permit counting of a uniform
suspension without the problems as-
sociated with cells settling while being
counted. A standard solution was pre-
pared by adding a fraction of a milliliter
of the same solution injected into the
animal to saline in a total volume of 1
liter. The standard was prepared for
counting by the same procedure as the
blood specimens.
All samples were counted for 100
minutes in a 3-inch well Til crystal
attached to a Packard Model 2001
Spectrometer Sealer-timer. Counts of
the paired blood specimens were very
close, within 5 percent in December
and 3 percent in March. The reported
blood volumes are the mean values
of the respective paired samples. The
oxyhemoglobin dissociation curve was
determined on 21 January 1972. A
20 ml blood sample was secured from
a puncture of a distal brachial vein
the the pectoral hn. The blood was
immediately placed into well-heparin-
ized (250 units heparin per 10 ml
blood) plastic test tubes, inverted 4 or
5 times, and then placed in an ice bath.
Less than 4 hours after collection
the dissociation curve was completed
by a Dissociation Curve Analyzer
(DCA-1. Radiometer, Copenhagen).
Duvelleroy et al. (1970) have ex-
plained in detail the methodologs in-
volved in the operation and construc-
tion of the 02-Hb dissociation curves
by the DCA-1. Slight changes in pH
were monitored for every point on
the dissociation curve. Because Pqo
changes with pH variation, corrections
were made to a pH of 7.4 with the
equation
AlogPp^
ApH
Bohr effect.
The value for the Bohr effect was
obtained from Lenfant ( 1969).
The hematocrit (Hct) was obtained
in the usual manner. A Clay-Adams
Autocrit Centrifuge was the instrument
used.
RESULTS
The hematocrit, blood volume, and
Pjo of the E. robitsius as well as cer-
tain physiological blood parameters
of other cetaceans are presented in
Table 1. The blood volume was deter-
mined on two occasions and the oxy-
gen binding capacity was determined
with one blood sample. Typical sig-
moid Oo-Hb dissociation curves are
shown in Figure 1. Curves for two
other cetaceans. Glohicephalci scani-
luoni and Tiirsiops i;illi, determined
by the same methods (Antonelis. 1972)-'
are included.
DISCUSSION
The hematocrit of Gigi is not un-
like that measured in other mammals.
While Lenfant (1969) asserts that this
is true for all marine mammals. Ridg-
way and Johnston (1966). Horvath et
al. (1968). and Ridgway et al. (1970),
have demonstrated an increased packed
cell volume in some of the small
cetacea. Lenfant (1969) attributes
such results to differences in technique
or in physical condition. However,
the animals who showed high hemat-
ocrits were maintained in captivity.
Had they followed the normal pat-
^ Antonelis, G A. 1972 Oj-Hb dissociation
curves of the pilot whale. Globicephala scam-
mom, and Pacific bottlenose porpoise, Tursiops
gilti- (Unpubl manuscr,}
tern of captive animals, the values
would have been even higher if sampled
in their natural environment. It has
been amply demonstrated that animals
brought into a captive situation soon
show a reduction in both hetnatocrit
and hemoglobin content (Gilmartin
and Ridgway, 1969,^* Lenfant, 1969).
The first blood volume for a large
cetacean using isotopic methods is
reported. Although H^i labeled hu-
man serum albumin was used in the
analysis, the similarity of the paired
blood specimens taken at each test
date indicates not only that mixing
was complete, but also that this foreign
protein was not being eliminated so
rapidly that a meaningful blood vol-
ume determination could not be made.
Unpublished data on the killer whale
are included also. Both animals have
blood volumes (£. rohusuis: 6.1 and
8.1 percent; O. onu: 8.2 percent)
within the range reported for other
species of large cetaceans; Laurie
(1933) reported a large blue whale's
blood volume as 6.6 percent. Smith
and Pace (1971) estimate that the
blood and body fluids of large ceta-
ceans to be between 10 and 15 percent
of the body mass.
Lawson { 1962) and Sjiistrand ( 1953,
1962) have reviewed the many factors
which affect blood volume and one
should be aware of them when evaluat-
ing blood volume data. Since the
animal is placed under highly stressful
conditions as well as the imposition
of unaccustomed gravitational forces
as a result of removal from the water,
the picture for marine mammals is
complicated.
Nutrition and electrolyte balance
also affect blood volume. To our
knowledge neither the freezing point
depression nor the osmolality of the
urine were determined. Osmolality
can be calculated, however, using the
formulas of Wolf (1958). Gigi's ex-
clusive squid diet must have produced
a urine whose minimum osmotic con-
3 Gilmartin, W G, and S H Ridgway 1969
Some physiological properties of the blood of
the killer whale, Orcinus orca- (Unpubl,
manuscr )
29
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en en
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in t
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o CNj -a-
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C\J O CT>
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eo
■-- CO 00
m in -^
r- en
in in
»- c\j m oj
x: T3
5-5
S 5
E S
a c
05 Q.
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o c
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E 5
o m
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— o
TO C
o —
£ ■?
(I) g
nr to 1^
TO £
5 CO
O) >
E o ^
III;
c 5 ii o - 5^ ;
ccntration was 1.670 milliOsniols/
liter. The effect of such a diet, which
is isoosmotic with sea water, is un-
known.
The P50, the partial pressure (mm
Hg) of oxygen at which hemoglobin is
50 percent saturated, is a measure of
the blood's affinity for oxygen — the
higher the P-m. the lower the affinity.
The P50 for Gigi was 36.5 mm Hg and
is the highest reported for any
cetacean. In a list of fifty mammalian
species in which the ^50 has been deter-
mined, only four animals exhibited a
higher P50 (Bartels, 1971).
Horvath et al. (1968) compared the
dissociation curves of several small
cetacea and found that a shift to the
left, or increased affinity for oxygen,
relates to individual species behavior
and feeding habits. Apparently this
pattern does not hold for the larger
cetaceans.
In looking at such parameters as
hematocrit, oxygen capacity, and par-
ticularly the P^o- a pattern emerges
and one is tempted to ascribe this to
some behavioral characteristic of the
animal. However, one should proceed
with caution.
Oxygen dissociation curves are de-
termined by several methods. The
method used by Horvath et al. ( l'^^68)
was to treat the blood with varying
levels of oxygen and sufficient CO2
to maintain a pH of 7.4. I, enfant ( 1969)
and .Schmidt-Neilsen and Larimer
(1958) maintained a PcO'? '^'f -*•' rnm
Hg in determination of their curves.
■Schmidt-Neilsen and Larimer (1958)
observed that In terrestrial animals
the blood of the larger animals has
the higher affinity for oxygen (low
P.w). Lenfant (1969) pointed out that
this does not hold for marine mammals
and the calculated Pcm's using their
formula supports this.
.Steen (1971) has pointed out that a
Pco2 ''' "^0 "I'll Hg may not repre-
sent the true arterial Pcoo '-^^ ^^'^
animal, e.g., a cat is about 28 mm Hg.
.Since the ^002 ^^as a profound effect
on the Pm and the magnitude of the
Bohr shift, one is hard put to make
meanlnglul comparisons in marine
30
mammals when different methods arc
used. To compMcate the picture fur-
ther, Riggs (I'JftO). using a buffered
system at a pH of 7.4. observed that
animals of varying size had identical
Pso's at that pH.
In order to make meaningful evalu-
ations of the dissociation curves in
marine mammals, the in vivo ^002
and pH need to be determined. Rieu
and Hamar (1968) point out the dif-
ficulties of drawing a representative
arterial blood sample although these
arterial data have been collected from
one species. Tiirsiops iruncalus, by
Ridgway (1968). In short, more work
needs to be done.
ACKNOWLEDGMENTS
We thank k. Suwa and E. Wahren-
brock of the Anesthesia Department.
University Hospital, San Diego, Cali-
fornia, for making the Dissociation
Curve Analyzer available for our use.
and the staff of .Sea World for their
cooperation.
This study was supported in parts
by grants from the Oceanography
Section of the National Science Foun-
dation (Grant #GA-3I297) and the
Bureau of Medicine (Project #MF-
12524014).
Ridgway, S. H. 1968. The botllcniise dolphin
in biomedical research. In W. I. Guy
(editor). Methods of animal experimenta-
tion. Vol. .^. p. 387-446. Academic Press,
New York.
Ridgway, S, H,, and D, G. Johnston. 1966.
Blood oxygen and ecology of porpoises
of ihree genera. Science (Wash., D.C.)
15I:456-4.S8.
Ridgwav, S. H., J. G. Simpson, G. S. Patton.
and W. G. Gilmartin. 1970. Hemato-
logic findings in certain small cetaceans.
J. Am. Vet. Med. Assoc. l.';7:S66-575.
Rieu, M„ and M. Hamar. 1968. Prelimi-
nary results concerning the gas oT arterial
and veinous blood in Dcli'luiui.\ dclphu
and Sli'iR'llu .vma. Report for ONR,
Contract NF6 10527 l)006.V
Riggs, A. 1960. The nature and significance
of the Bohr effect in mammalian hemo-
globins. J. Gen. Physiol. 43:7,37-752.
Schmidt-Neilsen, K... and J. L. Larimer.
1958. Oxygen dissociation curves of
mammalian blood in relation to body size.
Am. J. Physiol. 195:424-428.
Sjostrand, T. 1953. Volume and distribu-
tion of blood and their significance in
regulating the circulation. Phvsiol. Rev.
33:202-228.
. 1962. Blood volume. In W. F.
Hamilton (editor). Handbook of physiol-
ogy. Circulation. Sect. 2, Vol. 1, p. 51-62.
Am. Phvsiol. Soc, Wash., D.C.
Smith. A. H.. and N. Pace. 1971. Differ-
ential component and organ size rela-
tionships among whales. Environ. Physiol.
1:122-136.
Sleen, J. B. 1971. Comparative physiology
of respiratory mechanisms. Academic
Press, Lond.. p. 109.112.
Wolf, A. V. 1958. Thirst - physiology of
the urge to drink and problems of water
lack. Charles C. Thomas, Springfield,
p. 344-355.
MFR Paper 1052. From Marine Fisheries Review, Vol.
36, No. 4. April 1974. Copies of this paper. In limited
numbers, are available from D83. Tec finical Information
Division, Environmental Science Information Center,
NOAA. Washington. DC 20235.
MFR PAPER 1053
Feeding of a Captive Gray Whale,
Eschrichtius robustus
G. CARLETON RAY and WILLIAM E. SCHEVILL
LITERATURE CITED
Bartels, H. 1971. Blood oxygen dissocia-
tion curves: mammals. In P. L. Altman
and D. S. Dittmer (editors). Respiration
and circulation, p. 196-197. Biological
Handbook. Fed. Am. Soc. Exp. Biol.,
Bcthescda.
Bernharl. F. W., and L. Skeggs. 1943. The
iron content of crystalline human hemo-
globin. J. Biol. Chem. 147: 19-22.
Duvelleroy, M. A., R. G. Buckles, S. Rosen-
kaimer. C Tung, and M. B. I. aver. 1970.
An oxyhemoglobin dissociation analyzer.
J. Appl. Phvsiol. 28:227-233.
Horvalh. S. M.. H. Chiodi. S. H. Ridgway.
and S. Azar, Jr. 1968. Respiratory and
electrophoretic characteristics of hemo-
globin of porpoises and sea lion. Comp.
Biochem. Physiol. 24: 1027-1033.
Laurie. A. H. 1933. Some aspects of res-
piratKtn in blue and fin whales. Discov-
ery Rep. 7:363-406.
Lawson. H. C. 1962. The volume of
blood — a critical examination of methods
for its measurement. In W. F. Hamilton
(editor). Handbook of phvsiologv. Circu-
lation. Sect. 2, Vol. 1, p. 23-49. Am.
Physiol.Soc, Wash., D.C.
Lenfant, C. 1969. Physiological properties
of blood. In H. T. Andersen (editor).
The biology of marine mammals, p. 95-
116. Academic Press. New York.
ABSTRACT
Ihc fcc'diiii; of 11 captive yi'iirlini.; ft'iiuilc Eschrichtius robustus wirs oliscrvcd
wliilc (II villi; with Iwr «.v well as from llw surface. Slie sucked food off the lioliom
wliile \wiinniiiii; lipped over ahinil 120" so ilnil her c /uiA Ui/\ nearly parallel
to the lioUoni. .An increase in inoulli viiluine is apparently caused by action
of llie toni;ue, resulliui; in sironi; suction, dnriny; wliich ilie lower lip is opened
and food enters tlie inoiit/i. How food is separated from water and mud or
detritus is not l^nown. Tlie idiserved f>eluivior is protial^ly natural and illuminates
earlier records oj sionnuli cinuents. e.\teriuil inarl\iiis;s. and asyninielrii id
haleeii. Clearly, iniuli needs to he learned ahout tlie niecluuiisin of feeding; of
hideeii wluiles. I'liis species' feediiii; liahits nitiy he unic/ue anioiii; tliein.
INTRODUCTION
Observations on the food of Escli-
rielitius rohustus (Lilljeborg, 1861),
the gray whale, have, been summar-
ized by Zimushko and Lenskaya ( 1970)
and Rice and Wolman ( 1 97 1 ), indicat-
ing that the diet consists predom-
inantly of benthic animals, moslK am-
phipods and a few other crustaceans;
incidental items include polychaete
worm tubes, shells, gastropod opercula,
feathers, kelp, bits of wood. sand,
mud. and gravel. Tomilin (19.^7, p.
-346-347) suueests that Eschrichtius
r o
o o
~ 0)
o a
* °
0) z
._■ o
B E
2S
o S
32
G. Carleton Ray is with The
Johns Hopkins Univenity, Bal-
timore, MD 21205. His" work
was partly supported by con-
tracts to the University from the
Office of Naval Research (Oce-
anic Biologv). contract Nonr
N00014-67-A-0I63-0010 03.
William E. Schevill is with the
Museum of Comparative Zool-
ogy, Harvard University, and
the Woods Hole Oceanographic
Institution, Woods Hole, MA
02543. This is Contribution No.
3069 from the Woods Hole
Oceanographic Institution. His
work was partly supported by
the Office of Naval Research
(Oceanic Biologv), contract
Nonr N00014-66-()241.
is to some extent vegetarian (which
would make it a most exceptional
cetacean), supposing that seaweed
found in the stomach is food and not
merely incidentally swallowed. Howell
and Huey (1930) found the planktonic
Eiiplnuisiii pacificu in the baleen of
a gray whale taken off northern Cali-
fornia on 21 July 1926. Gilmore
(1961. p. 11) gives a winter observa-
tion of presumed feeding by gray
whales, '"criss-crossing thru a dense
school of small fish, like anchovies,
off San Diego," and Ken Balcomb
(pers. comm.) informs us that a gray
whale beached 15 miles north of
Grays Harbor. Wash., in April, had
a gullet packed with several gallons
of OsnuTus nionla.x (rainbow smelt).
It would appear that EsclvicltiiKs is
not limited to eating small benthic
crustaceans, but will also eat a variety
of other food as opportunity offers.
Gray whales seem to do most of
their feeding in the Bering and
Chukchi Seas (Pike. 1962, p. 831-8.32).
Rice and Wolman (1971. p. 24-2,S)
conclude that all organisms found
in the stomachs of gray whales killed
on the Arctic summer grounds are
"infaunal benthic species." They state
that 95 percent of the food species
found in one Bering Sea sample were
gammaridean aniphipods 6 to 25 mm
long, and that the predominant spe-
cies from this sample, Anipdiscu
iiuicroi cplnild. "occurs mainly en
sandy bottoms at depths of 5 to 300
meters" (italics oursl. Zimushko and
Lenskaya (1970) say that gray whales
feed on nectobenthos. some 70 spe-
cies in all. but that only six species of
amphipods are of primary importance.
We assume that though such active
creatures as amphipods may be in-
faunal at times, their well-known ten-
dency, when disturbed, to move just
off the bottom, would make them
readily available to a sweeping whale.
One of us (Ray) observed this amphi-
pod behavior from a submersible in
the Bering Sea in 1972. Nemoto
( 1959) has discussed probable feeding
behavior of whales in the light of
mouth shape and baleen characteris-
tics. During the last century many
authors have alluded to gray whales
surfacing with mud visible on the
beak or other dorso-lateral parts. Pike
(1962. p. 823) cites a particularly
illuminating communication from Dr.
F. H. Fay, who mentions a gray whale
supposed to be feeding in 5 fathoms:
"As this whale surfaced close to the
vessel, mud was seen washing from
its back." Although Tomilin (1957.
p. 347) supposes that these whales
may actually dig their mouths into
the bottom, scooping and plowing, it
seems to us (see below) that their
behavior and anatomy are better
adapted to sweeping the bottom than
for digging; this accords well with
the evidence of asymmetrical barnacle
infestation and baleen wear reported
by Kasuya and Rice ( 1970).
The gray whale calf. Gigi, which
was captured in March 197 1 and
kept by Sea World. San Diego, was
initially fed an artilicial diet, but was
soon taught to eat full-grown squid.
Lolii^o npulesccns. By the time of our
observations (28 January- 1 February
and 1 1 March 1972). her daily diet
was 900 kg of squid, dropped frozen
into her tank in 9 kg blocks, and her
weight gain was almost 40 kg a day.
Our behavioral observations were
made both from the water's surface
and by scuba-diving with Gigi. We
also used underwater motion pictures
made h\ John .Seeker of Sea World
when Gigi was about 6 months old.
To aid in our interpretation, we have
consulted her trainers (Bud Donahoo
and Susan Bailey), and we have solicit-
ed observations on Gigi and on other
gray whales from several of our col-
leagues. To them, who are mentioned
below, to the Naval Undersea Center,
and to the management of Sea World,
we are grateful.
BEHAVIORAL OBSERVATIONS
Before detailing our observations,
we remind the reader that a gray
whale's head is roughly triangular in
cross section, the gular region being
the base while the cheeks form the
sides, sloping inwards at about 60°
towards the narrow beak-like upper
jaw. The curved mouth is at about
the middle of the cheeks. The mouth
has effectively only lower lips; the
upper lips are represented by the
rabbet-like recess above the gum line
into which the lower lips fit snugly.
In the course of teaching Gigi to
eat squid, trainers Donahoo and Bai-
ley taught her to relax the edge of the
left lower lip and turn it outward in
response to light taps on the head.
Food was placed by hand in the open-
ing thus created, passing into the
throat either through or under the
baleen. This was in contrast to the
feeding of the artificial liquid diet,
when the jaws were opened while
accepting the feeding tube; during the
hand-feeding of squid, the jaws re-
mained closed and the lip was opened.
as was also the case later on when one
of us thrust an arm down her pharynx.
Training Gigi to move her lip volun-
tarily was critical, for normally the
lip was held so tightly shut that a man
could not forcibly pry it open. This
training was done while she was
grounded in the tank almost empty
of water; it was not long before she
would thus accept food while swim-
ming. Soon thereafter she was feeding
freely without the aid of her trainers.
Gigi was always fed from the left side
(Donahoo has mentioned having been
i2>
34
o
5 *
in n
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a o
35
o n
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tg
n
36
a horsLMiian before hecoming acquaint-
ed with whales), which may account
tor the left-sided sweeping behavior
described below; Kasuya and Rice
(1970) reported that of 34 gray
whales that they investigated, 3 1 were
right-sided.
We observed that the edges of the
lips could be turned out and down
through about 60°, either one or
both sides at a time. Motion pictures
further show a fluttering of the pos-
terior part of this edge during hand-
feeding, especially near the major
flexure.
Voluntar\ feeding was as follows:
The frozen blocks of squid floated at
the surface and, as they thawed,
the squid mostly sank slowly to the
bottom. Gigi often "nibbled"' at the
thawing corners of the blocks, using
the left side of her mouth and usually,
but not always, holding the block at
about the place where the fluttering
had been noted. The nibbling was oc-
casionally accompanied by a noisy
pulsation called "earthquaking" by the
trainers, and splashing or jetting of
water and air for nearly half a meter
(Figures 1 and 2). The jet was usually
at this same place near the after end of
the mouth, but it sometimes ran nearly
the entire length of the baleen (not
quite to the forward end of the mouth).
Often the jetting was on both sides
of the mouth. When Gigi's mouth was
at the surface, air was involved in the
jetting, but not always when her
mouth was completely submerged, and
not at all when she was on the bottom
of the tank. We assume that this air
was adventitiously taken in. as in
eating soup.
After most of the squid had fallen
to the bottom of the tank. Gigi's be-
havior altered markedly. As she ap-
proached them, she would roll over
toward her back some 120°, so that
her cheek was nearly parallel to the
bottom and about 10-20 cm above it.
As she swam over the squid, she left
a clean swath 30-50 cm wide. It was
apparent that the squid were being
sucked up in a sort of pulsation, as
some squid briefly reappeared after
their first disappearance into her
mouth. It is presumed that she could
easily see the squid lying in her path.
In the cylindrical tank she described
a track slightly dorsad of straight
ahead, so that she swept over the squid
at about a 30° angle to the mouth
(Figures 3 and 4). Then three separate
actions were seen: ( 1 ) an opening of
the edge of the mid to posterior part
of the left lip so as to fold it away
from the baleen. (2) a swelling of the
gular region and expansion of the
gular grooves (Figure 4). and (3) an
opening of the right side of the mouth,
during which squid were sometimes
jetted out. The third item may merely
mean that it is easier to open both
sides of the mouth symmetrically,
though Gigi had showed us that she
could flex her lips one side at a time.
Since we could not see all these parts
of the whale at once, we can only infer
the presumable sequence of these re-
lated events. Then Gigi righted herself
and swam away; sometimes turbid
jets could be seen pulsing from both
sides of her mouth.
ANATOMICAL
INTERPRETATION
Our understanding of the mecha-
nisms involved is hampered by our
ignorance of the myology and other
soft anatomy of this species. We have
been able to find only osteological
anatomical descriptions and have had
no carcass available for even rough
dissection. W. C. Cummings and J.
Sweeney made, on our account, some
exploratory sections of the lower lips
of dead neonates found on the beaches
of Laguna Ojo de Liebre, Baja Cali-
fornia, and found them to be well
muscled. D. W. Rice reminds us that
the tongue is also well muscled, much
more so than in Btilucnopicni; it is
well figured and described by Andrews
(1914, p. 254. pi. 2 1. fig. 4. and pi. 22,
fig. 6).
All this, as well as observations of
behavior, strongly indicates that the
gray whale's oral anatomy is adapted
for suction and thai motion of the
lips is voluntary. We had but limited
opportunity to manipulate Gigi's
mouth ourselves; one of us (Schevill)
had his arm in her mouth several times
while she was "earthquaking" and
could feel no motion at all of the
tongue and only a slight agitation
near the larynx. But W. E. Evans (in
litt.) states that "the tongue cannot be
pulled back and forth very easily;
however, it can be raised high, dis-
placing a reasonably good percentage
of the volume of the mouth cavity".
Donahoo had his hand in Gigi's mouth
repeatedly; both he and Evans have
emphasized the tongue's strength and
mobility. Donahoo asserts that it
moves so as virtually to vacate the
oral cavity and that this involves a
shape change. He further asserts that
the shape change travels rearward
and that this movement of the "ball"
of the tongue can be seen from outside,
as the gular grooves expand. This
posteriorly moving expansion of the
gular region was also seen by one of
us (Ray) underwater. Further, Dona-
hoo said that as the tongue moves back,
a strong inflow appears at the out-
folded lip. He added that Gigis
feeding was not simply accepting,
but quite selective. When presented
a mixture of squid, "Pacific mackerel "
(chub mackerel, Scomhcr juponiciis).
and "whitebait" (probably jacksmelt.
Atlu'iinopsi\ culiforiiiciisis. or top-
smelt. Alhcrinops affinis). all three
were sucked from the bottom, but
only squid were retained, the others
being rejected.
CONCLUSIONS
Nothing benthic of the size of squid
has been reported in the diet of
Eschiichlius. so we should be cautious
in interpreting this captive's feeding
style as indicative of natural behavior
of the species, bearing in mind that
Gigi was completely isolated from her
kind throughout captivity. Neverthe-
less, her bottom-sweeping habit we
suppose may be natural, since it ap-
pears appropriate for catching the
animals that comprise the recorded
37
natural food of this species of whale.
Our captive's habit of sweeping a few
centimeters off the smooth tank bot-
tom does not deny the probabilits
that sweeping a soft or irregular
bottom at sea could get mud on the
sweeper's back (cf. Fay in Pike, 1962.
p. 823), especially if the prey is
actually benthic.
Cetological literature is full of poor-
ly supported conjecture, and we hesi-
tate to add more. Although we have
learned a number of things from the
captive Gigi, there is still much un-
known. For one thing, her jetting
water in pulses from a particular re-
stricted part of her mouth seems to
imply, perhaps, a special activity of
the tongue. Furthermore, we do not
understand the mechanics of the hy-
draulics that bring the food-bearing
water into the mouth. This is no mys-
tery in whales that swim along with
the mouth wide open, but it is not so
obvious in a whale which swims along
rather slowly with only a narrow slit
open, as did our Esclirichiius. Here it
seems necessary to increase the volume
of the mouth to cause useful inflow of
water. We are handicapped by our
imperfect understanding of the func-
tions of the muscular tongue. W. E.
Evans (pers. comm.) has told us that
Gigi's tongue once pressed his hand
painfully hard against her palate. Such
pressure might serve to push the gular
region downward, enlarging the mouth
cavity, and this idea fits with the ob-
servations of Donahoo and Ray of
the migrating tongue-bulge visible
from beneath.
Thus we suppose, from the assorted
evidence, the following concatenation
of events in feeding; First the whale
rolls over far enough so that the cheek
is about parallel with the bottom, and
the lip is opened as the tongue, press-
ing against the palate, pushes the gular
region away so that it expands, pro-
ducing an inflow which brings in the
epibenthic food. Then the tongue
rcla.xes and the gular musculature
tightens, reducing the size of the
mouth cavity and expelling water;
the food IS trapped in the baleen
fringes. We do not know exactl> what
happens next; perhaps a slight re-
newed suction of water removes the
food from the baleen fringes, and
swallowing presumably follows.
LITERATURE CITED
Andrews. R. C. 1^*14, Monographs of the
Pacific cetacea. I. The California gray
whale {RInuhtanccU's tiUmcita Cope).
Mem. Am. Mus. Nat. Hist. (New Series)
1:2:7-287.
Gilmore, R. M. 1961. The story of the
gray whale. 2ncJ ed. Privately published.
San Diego. 17 p.
Howell, A. B., and L. M. Huey. 1930. Food
of the gray and other whales. J. Mammal.
11:321-322.
Kasuya, T., and D. W. Rice. 1970. Notes
on baleen plates and on arrangement of
parasitic barnacles of gray whale. Sci.
Rep. Whales Res. Inst. 22:39-43.
Nemoto, T. 19?9. Food of baleen whales
with reference to whale movements. Sci.
Rep. Whales Res. Inst. 14: 149-290.
Pike. G. C. 1962. Migration and feeding
of the gray whale {E.\chruhiiiis nihhusiis).
J. Fish. Res. Board Can. 19:815-838.
Rice. D. W., and A. A. Wolman. 1971.
The life history and ecology of the gray
whale (E.schnchtius nihusius). Am. Soc.
Mammal.. Spec. Publ. No. 3. 142 p., 18
tables. 38 text figs.
Tomilin, A. G. 1957. Kitoobraznye. Zveri
SSSR i prilezhashchikh stran (Cetacea.
Mammals of the USSR and adjacent coun-
tries.) 9. 756 p. (Engl, transl. Smithson.
Inst., 1967.9,717 p.)
Zimushko. V. V., and S. A. Lenskaya. 1970.
O pitanii serogo kila (Eschrichinis gib-
hosiis Erx.) na mestakh nagula (Feeding
of the gray whale [E.schnclunis nihhosiis
Erx.) at foraging grounds). Fkologiya
Akad. Nauk SSSR l(3):26-35. (Engl,
transl.. Consultants Bureau. Plenum Publ.
Corp., 1971. Fkologiya 1(3):205-212.)
MFR Paper 1053. From Marine Fisheries Review, Vol.
36, No. 4, April 1974. Copies of this paper, in limited
numbers, are available from D83. Technical Information
Division. Environmental Science Information Center.
NCAA. Washington. DC 20235.
MFR PAPER 1054
Sounds Produced by the Gray Whale,
Eschrichtius robustus
JAMES F. FISH, JAMES L SUMICH, antj GEORGE L. LINGLE
ABSTRACT
U ndcrwawr \iiuihIs pnuliKcil hy a yniiiii; cupiivc yn/y uluilc arc described.
A "iucudUc-\t>it}idiiii> pulsed signal." consisliiiii of H lo 14 pulses in hursts Uisliui;
up Id 2 sec was the niosl c<>nuiu>n vocalization. Other souitds included a low-
frequency "i;ro\\l" or "moan." similar lo a sound recorded from L;ray w/iales at
sea: a short, hroiulhand. "i;riinllike" sound: a low-piit hed "hlowluile rumble":
aiul a loui; "ntctaUic-souiuliui; pulse train" tlial meiiicd into a low-frequency
"Kroan." The sounds ctnild not be correlaled with specific behaviors. Also de-
scribed are "clicks" recorded in the presence of the whale wlien site was returned
to \ea ami similar "iliiks" recorded from i;ray whales in liii kaninnish Bay.
I'ancoin er Island. Caitoila.
This report describes a variets of
sounds recorded from Gigi, a young
gray whale. Eschrichtius robustus.
while she was in captivity at Sea
World, a marine park in San Diego,
Calif., and sounds recorded in the
vicinits of the whale when she was
returned to the ocean nearly a year
later. Also described are the sounds
recorded in the presence of gray
whales in Wickaninnish Bay, Van-
couver Island. Canada.
38
Table 1. — Summary o( previously published data on gray whale sounds.
Eberhardt
& Evans.
1962
Painter. 1963
iWenz. 1964
Rasmussen & Head. 1965
2Gales. 1966
Hubbs, 1966
Asa-Donan & Perkins. 1967
Cummings et al , 1968
Poulter. 1968
Pulse
Repetition
Pulse
Peak
Duration
Frequency
Pulses per
rate
duration
energy
Signal type
(sec)
(Hz)
burst
(per sec)
(msec)
(Hz)
"Croaker-hke
40-700
80-300
grunts"
■Low-frequency
40-700
80-300
rumbles '
"Pulses"
4-6
100
"Low-pitched
4-9 pulses
Approx 33
11
grunting"
per grunt
"Clicks"
<200->3.000
10
No sounds
"Clicks-
<200->3,000
10
No sounds
"Ectiolocation-
70-3.000
5-22
3-7
1-15
400-800
like pulses"
"Variable
whistles"
"Moans"
1 5
20-200
"Underwater
Approx. 1
15-175
<1C0
blow"
"Bubble-type
0 7
15-305
sounds"
"Knocks"
to 350
"Croak-like
grunts"
"Rumbles"
"Cries"
"Grunting"
"Rasping"
1 to several
"Pulses"
5-18
"Chirps"
2-5
8
"Bong"
"Clicks"
to 12,000
3-5
'■- Both references show data on sounds recorded by Asa-Donan in 1955
Vocalizations have been recorded
from migrating gray wiiales off the
southern California coast (Wenz, 1964.
and Gales. 1966. both reporting on
recordings made by P. V. Asa-Dorian
in 1955; Asa-Dorian and Perkins.
1967: Cummings. Thompson, and
Cook. 196S) and from gray whales
in the lagoons of Baja California.
Mexico, where the whales breed
(Eberhardt and Evans. 1962; Painter.
1963; Poulter. 1968). Unsuccessful
attempts to obtain sounds from gray
whales off southern California and
in the lagoons have been made by
Rasmussen and Head (1965) and by
Hubbs (1966). The published data
on gray whale sounds are summarized
here in Table 1 .
Gigi had already been in captivity
at Sea World and hand-fed by her
trainer for about 2 months before the
tank recordings were made. Although
she seemed quite content in her un-
natural surroundings, her behavior
was certainly not representative of a
free-ranging gray whale of the same
age. Hence, the sounds may or may
not be similar to sounds emitted by a
young gra\ whale in its natural en-
vironment. A second problem with
any tank recording is the effect of
tank resonance and reverberation on
the physical characteristics of the
sounds. Certain frequencies were
probably accentuated in amplitude
and extended in time. Nevertheless,
the data at least represent the general
James F. Fish is with the Naval
Undersea Center, San Diego,
CA 92132. James L. Sumich is
with Grossmont College, El
Cajon, CA 92020. George L.
Lingle is with SEACO, Inc. al
the Naval Undersea Center, San
Diego, CA 92132.
frequency range and variety of a
young captive gray whales sound
emissions.
SOUNDS OF GIGI
AT SEA WORLD
Sounds were recorded simultane-
ously in water and in air on a 2-track
tape recorder (Uher 4200)' at 19
cm/sec. The hydrophone (Wilcoxon
M-H90-A). connnected to one chan-
nel of the recorder, was suspended 1
m above the bottom of the circular
concrete tank (11 m wide X 4 m
deep). The frequency response of the
underwater recording system was 40
Hz to 16 kHz. ±.^ dB. The micro-
phone, connected to the other channel.
was lowered over the lip of the tank
' Use of trade names in this publication does
not imply endorsement of commercial products
by the National Marine Fisheries Service.
39
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to record the commentary of the
trainer in the tank with the whale.
The 3-hr recording session began
about I hour before a feeding period
and lasted until the water level, which
initially was about I m above the
whales back, was too low to make
useful underwater recordings.
■Speclrographic analyses were made
in the laboratory with a "Vibralyzer"
(Kay Electric Company) to determine
frequency vs. time, and a real-time
spectrum analyzer (Spectral Dynam-
ics SD330) connected to an X-Y re-
corder (Hewlett Packard 7035B) to
portray the relative amplitude vs.
frequency. The waveforms were moni-
tored with either the spectrum an-
alyzer in the Scope Time mode or an
external oscilloscope. All of the
sounds described below were recorded
from the hydrophone output.
The whale was very inactive and
emitted no sounds until the water was
lowered enough for the trainer to
stand in the tank and touch her back.
None of the sounds could be consist-
ently associated with a particular
behavior. However, one type, a "me-
tallic-sounding pulsed signal." was
emitted nearly every time the trainer
tapped the whale lightly on the back.
A low-frequency "growl" or "moan."
similar to one type of sound recorded
from gray whales off San Diego.
Calif., by Cummings et al. (1968).
was produced twice during the re-
cording session. The principal energy
of this signal recorded from the cap-
tive animal was in a band from 100
to 200 Hz, with a secondary peak
around 1.5 kHz (Figure lA). The
duration of the sound was just over 1
sec. There was no obvious movement
of the blowholes or expulsion of air
associated with this vocalization.
The most common sound was the
"metallic-sounding pulsed signal"
which consisted of 8 to 14 pulses in
bursts lasting up to 2 sec (Figure IB).
The pulses had sharp fronts (fast
rise times) with energy extending from
below 100 Hz to over 10 kHz. and
several resonant peaks, the strongest
being at 1.4 kHz. This sound occurred
as often as five times a minute, even
when not incited by the trainer. Only
occasionally did it appear to be cor-
related with exhalation and move-
ment of the blowholes.
Three times during the recording
session, a short (0.2 sec), broadband,
"gruntlike" sound (Figure IC) was
emitted, without movement of the
blowholes. Its peak energy was cen-
tered at 200-400 Hz and 1 .6 kHz.
Figure ID shows the underwater
sounds of an exhalation followed by
a low-pitched, "blowhole rumble."
This combination occurred several
times.
Twice, a long "metallic-sounding
pulse train" with a repetition rate of
about 14 pulses/sec merged into a
long, low-frequency "groan" after
about 1.5 sec (Figure IE). Except for
the much faster pulse repetition rale,
the first part of this vocalization was
similar to the sound shown in Figure
IB.
Numerous other sounds produced
by Gigi during the 3-hr recording
session essentially were variations of
one of the five types discussed above.
SOUNDS RECORDED DURING
RELEASE OF GIGI
Unfortunately, we did not record
again in the presence of Gigi until
she was released on 13 March 1972.
The recording and analysis system
used for these data was the same as
used at Sea World. Shortly after Gigi
was lowered into the water from the
barge that carried her out to sea,
long trains of "clicks" were heard.
Although at the time there was no
wa> to determine if these sounds,
which were unlike any recorded from
Gigi at Sea World, actually came
from the whale or from another un-
seen biological source in the area, we
now believe they were emitted by Gigi.
The clicks were nearly identical to the
clicks we have recently recorded in
the presence of gra\ whales in Wick-
aninnish Bay, Vancouver Island. Can-
ada.
The clicks recorded in the presence
of Gigi are shown in Figure 2. Their
principal energy occupied a band
from about 2 to 6 kHz, centered at
3.4 to 4.0 kHz. Click duration was I
to 2 msec. Eight minutes and 15 sec
after the whale entered the water
most boats in the area shut down their
engines for our recording. The first
burst of 29 clicks was recorded 6 sec
later. Three minutes and 49 sec later
the boats started their engines and we
had to terminate our final recording
of Gigi. During the 3 min and 55 sec
of quiet-ship conditions we recorded
1.304 clicks. The number of clicks
per burst (or train) varied from I to
833 and the click repetition rate from
9.5 to 36.0/sec. The longest click
train, containing 833 clicks at an
average repetition rate of 19/sec.
began about 1 min after the boats had
shut down their engines. Although
the amplitude of the signals varied
with time, we could not correlate
signal level with the location of Gigi
because the animal was not seen dur-
ing the entire time of the recording.
SOUNDS RECORDED FROM
GRAY WHALES OFF
VANCOUVER ISLAND
The system used to record sounds
in the presence of gray whales in
Wickaninnish Bay on the west coast
of Vancouver Island, Canada, con-
sisted of a cassette recorder (Sony
Model TC-126) and a portable under-
water listening set (InterOcean Model
9()A Bio-Acustik). The useable fre-
quency range of the system was 100
Hz to 10 kHz. The hydrophone ar-
rangement shown in Figure 3 resulted
in good quality recordings with the
small boat system.
Since 1967, as many as seven gray
whales have been sighted at one time
in Wickaninnish Bay. However, all of
the recordings described here were
from single whales or pairs. At 1725
hr on 10 August 1973. several click
trains were recorded from a single
feeding gray whale in 10 m of water,
1.200 m friim shore. Ver\ little wind
42
and calm seas made recording condi-
tions ideal. The first clicks, shown in
Figure 4F, began 1 mm after the
whale started a 3-min-J!5-sec-long dive.
at a distance of 50 to 70 m from the
hydrophone. Additional click trains
(Figure 4G) occurred simultaneously
with the first exhalation after the
dive. Twenty sec later, noise from an
unseen boat began and continued for
95 sec. A third click train was emit-
ted 50 sec after the boat noises ceased
and 50 sec prior to the next blow. By
then, the whale was 80 to 100 m from
the hydrophone and the received level
of the clicks was 5 to 7 dB lower than
the level of the clicks recorded when
the whale was half that distance from
the hydrophone.
On 18 August 197.^. the click train
shown in Figure 4H was recorded
from a single feeding gray whale at
0900 hr. The whale was about 600 m
from shore in 4 m of water. The sur-
face was calm with about a Im swell.
At the time the click train was emitted,
the whale v\as 100 to 150 m from the
hydrophone. Twenty min later a sin-
gle harbor porpoise, Phocoena pho-
coena. was observed in the area.
About 5 hr of recordings uere
made in the presence of the gray
whales in Wickaninnish Bay and
much additional monitoring was done
without recording. Although at times
nearly continuous very faint clicking
could be heard, only about 250 of the
recorded clicks had good signal-to-
noise ratios. The number of clicks per
train varied from I to 96 with repeti-
tion rates of 8 to 40/sec. The principal
energy of these clicks occupied a band
from about 2 to 6 kHz. centered at
3.5 to 4.0 kHz. The average click
duration was a little under 2 msec.
DISCUSSION
striction (similar to the sound of air'
escaping from a scuba regulator un-
derwater). Since this whale sound
generally was not associated with ex-
halation or blowhole movement, if it
were, in fact, generated by escaping
air. the air must have passed from one
internal chamber to another. No bub-
bles v\ere observed coming from the
mouth or blowholes.
Although the possibility exists that
another species of marine mammal
could have produced the clicks re-
corded when Gigi was released off
San Diego and the clicks recorded in
the presence of gray whales in Wick-
aninnish Bay. we think the evidence
indicates that the clicks did come from
the gray whales. The acoustic param-
eters of the clicks recorded from the
geographic areas are nearly identical.
The only marine mammals, other
than gray whales, observed in either
recording area was the single Phocoena
phocoeiw observed a half hour after
the recording was made on 18 August
1973 in Wickaninnish Bay and a
small group of Dclphiniis dclphis.
about 2 km awa\ from the site of
Gigi's release a half hour before she
was released. Phocoena phocoena.
however, has not been observed off
San Diego, and clicks of Delphinits
have a much higher frequency content
than described in this report. Also,
the level of the clicks recorded in the
presence of Gigi was too high for the
sounds to have come from the Del-
phinits as the clicks appeared to origi-
nate from a single source rather than
from a group of animals.
We have no evidence that the clicks
recorded in the presence of gray
whales have an echolocation function,
but if the\ do. their frequency range
(2 to 6 kHz) probably would be too
low for the sounds to be useful for
locating small individual food organ-
isms. However, they could be helpful
for finding dense concentrations of
organisms or for ranging off the bot-
tom to feed or navigate. Despite four
seasons of recording in the presence
of hundreds of migrating gray whales
off San Diego, Naval Undersea Center
personnel have never recorded similar
clicks from the whales. But. accord-
ing to most authorities, gray whales
do not feed on their long migrations
(Rice and Wolman. 1971). If the
clicks were associated with feeding,
we consequently should not expect to
TAPE RECORDER
AMPLIFIER
CORK FLOATS,^
HYDROPHONE LINE
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SPAR BUOV (BICYCLE
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BRASS SWIVEL
8" DAMPING DISC -
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NEUTRAL BUOYANCY
We do not know how any of the
sounds discussed in this paper were
actually produced by the gray whales.
The ""metallic-sounding pulsed signal"
produced by Gigi at Sea World sound-
ed like air bubbles escaping from an
area of high pressure through a con-
Fjgure 3. — Hydrophone suspension system used to record underwater sounds in Wickaninnish
Bay, Vancouver Island, Canada.
43
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encounter them in this area of migrat-
ing whales. When the clicks were
recorded in Wickaninnish Bay, the
gray whales were feeding. Why Gigi
emitted clicks when released is un-
known. In this case, their function
could have been orientation since it
is unlikely that she was looking for
food so soon after being placed in a
new environment. The clicks dis-
cussed here are only slightly like
those recorded by Asa-Dorian in 1955
(see Wenz. 1964). They are not similar
to any other reported gray whale
sounds.
Other recent evidence for mysticetes
producing click-type sounds has been
reported by Beamish and Mitchell
(19711. Their recordings in the pres-
ence of blue whales included clicks
with peak energy in a band from 2 1
to 3 1 kHz.
ACKNOWLEDGMENTS
We are grateful to Wilburn G.
("Bud") Donahoo for his help at Sea
World during the recording session;
to William C. Cummings for his
advice and assistance in the recordings
of Gigi at sea. and to William E.
Evans and Fay Wolfson for their
suggestions on the manuscript.
This work was supported by the
Naval Undersea Center. Independent
Research funding, and the Office of
Naval Research. Oceanic Biology
Branch. Grant No. NR 104-123.
LITERATURE CITED
Asa-Donan, P. V., and P. J. Perkins. 1967.
Ttie controversial production of sound
by the California gray whale, Eschricluiiu
gihhosus. Nor. Hvalfangst-Tid. (Norwe-
gian Whaling Gazette) 56:74-77.
Beamish, P., and E. Mitchell. 1971. Ultra-
sonic sounds recorded in the presence of
a blue whale, Bakwrniplera tnit^cultis.
Deep-Sea Res. 18:803-809.
Cummings, W. C, P. O. Thompson, and R.
Cook. 1968. Underwater sounds of mi-
grating gray whales, Eschruhtius kUiucus
(Cope). J. Acoust.Soc. Am.44:1278-1281.
Eberhardt, R. L., and W. E. Evans. 1962.
Sound activity of the California gray
whale, Eschrichnu.s glaiiciis. J. Aud. Eng.
Soc. 10:324-328.
Gales, R. S. 1966. Pickup, analysis, and
interpretation of underwater acoustic data.
Ill K. S. Norris (editor). Whales, dolphins,
and porpoises, p. 435-444. Univ. Calif.
Press, Berkeley and Los Angeles.
Hubbs, C. L. 1966. Comments. In K. S.
Norris (editor). Whales, dolphins, and
porpoises, p. 444. Univ. Calif. Press,
Berkeley and I os Angeles,
Painter, D. W., II. 1963. Ambient noise
in a coastal lagoon. J. Acoust. Soc. Am.
35:I458-1459(L).
Poulter, T, C. 1968. Vocalization of the
gray whales in Laguna Ojo de Liebre
(Scammon's Lagoon) Baja California,
Mexico. Nor Hvalfangst-Tid. (Norwegian
Whaling Gazette) 57:53-62.
Rasmussen, R. A., and N. E. Head. 1965.
The quiel gray whale ( E.schnchinis glaiicu.s).
Deep-Sea Res. 12:869-877.
Rice. I). W., and A. A. Wolman. 1971.
The life history and ecology of the gray
whale {Escliricluiiis robuxius). Spec.
Publ. 3. Am. Soc. Mammal., 142 p.
Wenz, G. M. 1964. Curious noises and
the sonic environment in (he ocean. In
W. N. Tavolga (editor). Marine bio-
acoustics, p. 101-123. Pergamon Press.
New York.
MFR Paper 1054. From Marine Fisheries Review, Vol.
36, No. 4, April 1974. Copies of this paper, in limited
numbers, are available from D83. Technical Information
Division, Environmental Science Information Center,
NCAA, Washington. DC 20235.
MFR PAPER 1055
Aerial Observations of Migrating Gray
Whales, Eschrichtius robustus, off
Southern California, 1969-72
J.S. LEATHERWOOD
ABSTRACT
Miiiratini; iiray hIuiIcs were observed from lielicopler und fi.\ed-\\ini; (linnifl
from central Ccdifoniici .soiilh lo Cedros mid Giuididiipe Islunds. Baja California.
Me.xico. Willi ihe primary .\ii;/ilini.; effort off southern California. Peak nundiers
were observed off soiit/iern California in January for tlie southward iniiiration
and in Marcli for tite iiortliward inii;ratioii. Individuals were observed with the
same relative frequency 80-160 km offslun-e as they were within 80 km of
shore. Cows witli calves were seen from February through May. primarily
inshore, and tended lo be alone or with other cou s with calves. Yearling whales
were seen insliore from February through April and also tended to be solitcuy
or with other yearlings. Average speed of movement for lunthward migrants
was 2.8 knilhonr.
Results of aerial surveys compare favorably with published summaries of
the liming of migration based on sliore and ship samples and support the \'alue
oj aerial surveys us u tool in cetacean population studies.
INTRODUCTION
Since shortly after its population
began to recover from a second near-
e.xtermination by man in the 1920's
and I930's (Gilmore, 1955), the Cali-
fornia gray whale, Esclirichtius ro-
bustus, has been the subject of more
public interest and more scientific
research than perhaps any other spe-
cies of large whale. Because of their
spectacular nature and proximity to
shore along much of the route, the
migrations of the species have been
rather exhaustively described by Scam-
mon (1874), Hubbs (1959), Gilmore
(1960a and 1960b), Rice (1961), Pike
(19621. Hubbs and Hubbs (1967).
Adams ( 1968). and Rice and Wolman
(1971). Observations from shore sta-
tions (primarily at Point Loma in
45
San Diego, and at Yankee Point near
Monterey) supplemented with aerial
observations and boat surveys, have
fixed the timing and described most
aspects of that migration in detail.
Even so. several interesting gaps
still exist in our knowledge of the mi-
grating animals. For instance, although
Gilmore (1969) has discussed move-
ment patterns of yearling whales on
the southern migration, there are no
reports on the movements of yearlings
during the northern migration. Simi-
larly, although Hubhs (1959) reported
that "cows with calves seem to take a
more offshore path." actual data on
-
_
-
I97Zn
-
1971^
19700
-
^
19S9n
-
'
-
GRAND TOTALS
-
-
74 FLIGHTS
279.70 HOURS
_J
— j
U 1 1— r-r
^
p-r^
!^^^x^4'
JAN FEB MM APR MAY JUN JUL AUG SEP OCT NOV DEC
Figure 1. — Flight hours, 1969-72.
the movements of mothers with calves
after they leave the breeding lagoons
are so scant that Rice and Wolman
(1971) simpl>' report that '"the route
taken by females and calves during
the spring migration is unknown."
Finally, although average rates of
movement for the population have
been computed from dates of peak
passage at two separate shore stations,
there are actual numbers for rates of
movement of individual animals to
test those averages only for the south-
ward migration (W\rick. 1954; Cum-
mings, Thompson, and Cook. 1968).
Since February of 1969. the author
has been conducting routine aerial
surveys of the cetaceans off southern
California, primarily in the area from
lat.34°N south to Islas Todos Santos
and offshore as far as 280 km.
Additional flights have surveyed the
coast from Point Mugu north to Mon-
terey Bay in March and the area
from San Diego to Cedros and Guada-
lupe Islands in January and February.
Ill 3^41(UP3 3 1 3^^,„„„
^-J 2 1 1 2 5lM 4 1 2 1^^
T|?1l 1 1 1 4 911 5 4 2 I'l'jJCEAN
1J2|1 IJ 2^^5 4 5 5 62" '™^
Wlfri 1 !i8 5 4 7 9 9 8
ijzil'l 1 2 2 3 412 62431*^
142 15 3 2 12 3 I02J3221
1 2
23322533366 y
3!2 2 2 1 1 14 2^
1 2 1111 1*1-|32'
Figure 2. — Survey effort by 10-minute blocks,
southern migration, 1 October to 15 February.
Each number represents total number of times
that area was surveyed during this time period.
METHODS
Flights were made in Navy H-3
helicopters and S-2 reconnaissance
aircraft, and in twin-engine rental air-
craft. Survey altitudes ranged from
150 to 300 meters depending on
weather conditions. Detailed observa-
tions were made from as low as 15
meters. Although gray whales were
not the exclusive target of the survey,
for every gra\ whale sighting the
number and estimated size of indi\id-
uals. their location, swimming direc-
tion and speed, and details of behav-
ior were recorded.
Through I July 1972 we made 74
flights totaling 279.7 hours of obser-
vation time (Figure 1). Sampling ef-
fort was accelerated during March.
April. May. and June 1972 in support
of the radio track of the gray whale
Gigi (Evans. 1972. and this publica-
tion) and of a common dolphin. Dcl-
pliiiui\ ilclphis. subsequently tagged
and radio-tracked from aircraft (Evans
and l.eatherwood. 1972). In general,
surveys were more extensive during
the period of the northern migration
(approximaieh mid-February through
Mav).
J. S. Leafherwood is a member
of the staff of the Bio-Systems
Program. Naval Undersea Cen-
ter, San Diego, CA 92132.
To facilitate data analysis, the
study area was divided into 10-minute
blocks, and tallies were maintained
of the number of times each zone was
surveyed for cetaceans whether or not
animals were sighted. Zones were not
recorded as surveyed if cloud cover,
fog. or surface water conditions pre-
vented adequate observation in the
area.
I
V
1
1 1; T^
r
s
^
^
1
>rT\ ^
1
112 2 iVa
1
2 2 2 2 4^ "
X
1
1
1
2
6 5 8 11J4 9 5 e"^
4 r,.». =T
1 2
1
1
1
1
14 4 3 131SiB9 10 7 7^7.
i |l
r 1 3 5 8 10171210 8 7
OCEAN
r2
I'l 1
1 2 1 5 f,\3 4 9 25131414 3, -|
1
l' :i'2 'l 1^4
8101631lt,„
1
4
4
5
5
6 5!8 4 8 14153139
s«
1
1
1
3
3
3
3^2(4 3 5 5 1015232^
1
1
2
2
2
3
5
V
2
2
1
2
4
4 3
3
3
2 1
2
3
1
1
1
1
1
-
1
1
1
1
1
1
liiii
1
1
»
1
1
1
1
1
\
1
12C
"
119"
HE
•
117
Figure 3, — Survey effort by 10-mtnute blocks,
northern migration, 16 February to 1 June, Each
number represents total number of times that
area was surveyed during this period.
Figures 2 and 3 summarize the to-
tal number of times each zone was
surveved during the periods of the
southern migration ( I October to 15
February) and northern migration (16
Februar> to I June) of gray whales.
Effort was concentrated off San Di-
ego during both periods because all
three airfields used are located there.
The substantial increase of effort in
the southern San Pedro Channel from
the southern to the northern migra-
tion is the result oi the aerial radio-
tracking mentioned above.
During the three migratory seasons,
gray whale groups were sighted 91
times in the stud> area. Of these. 23
groups included mothers with calves,
19 included yearling whales, and 8
sightings represented observations of
the same animals on successive davs.
46
RESULTS
Migration Peaks and
Offshore IVIovements
The picture of migration peaks ob-
tained from the aerial surveys agrees
with the summaries of Hubbs (1459)
and Gilmore (1960). The earliest
animals were seen in the third week
of December and the latest during
late May. The largest numbers of
animals were seen during the first and
second weeks of January and the sec-
ond and fourth weeks of March. Be-
cause the amount of aerial survey
effort varied from month to month,
indices of apparent abundance were
computed for data in blocks of a
month by dividing both the number
of aerial observations and the number
of individuals seen by the amount of
survey effort during that time period.
These indices (Figure 4) also clearly
indicate the periods of greatest abun-
dance off San Diego as January and
March.
During both legs of the migration
many whales were sighted far offshore.
(Figure 5) presumably taking what
has been called the "inter-island leg"
(Gilmore. 1969). For instance, with-
in the 64 km wide band between lat.
32°I5'N and 32°55'N. southern mi-
grating gray whales were encountered
during 4.7 percent of the flights in
the first 80 km from shore. 5.0 percent
of the flights in the second 80 km.
and 1 percent of the flights over the
next 48 km. Similarly on the northern
migration, grays were sighted 6.4.
9.2, and 3.3 percent respectively of
the times the three zones were sur-
veyed. These findings support the con-
tentions of Rice (1965) and Rice and
Wolman (197 1) that at least since
1965 a rather high percentage of the
whales have passed offshore, out of
sight of Point Loma.
Though most of the animals taking
the offshore route apparently strike
for the coast shortly after they pass
bv the southernmost of the Channel
5.0-
4.0
3.0
2.0
1.0
5.38
2.36
TOTAL DBS.
1" TOTAL FLT. HRS.
2.20
1.50
.38
.02
14.0
-
TOTAL ANS.
2 "TOTAL FLT. HRS.
12.6
12.0
-
10.0
-
8.0
-
6.1
6.0
—
4.91
4.0
"
2.4
2.0
"
.9
1.04
OCT NOV DEC JAN FEB MAR APR IMAY JUN OCT NOV DEC JAN FEB MAR APR MAY JUN
Figure 4. — Indices of apparent abundance (Ij) ol gray whales (rom aerial surveys. 1969-72.
34°
33 Figure 5. — Aerial survey
gray whale sightings,
1969-72.
32°
1?0°
119°
118°
117°
Islands, some do pass offshore toward
Guadalupe Island. Gilmore (1955) re-
ported Hubbs' sighting of three
mothers with calves outside Guadalupe
in February of 1950. In February
1972. I located two gray whales on
the outside of Guadalupe near the
southwest tip. A third animal, too
close to the cliffs to permit close ex-
amination, was also believed to be a
gray. The two verified sightings were
both adult animals 1 1 or more meters
in length.
Cow-Calf Groups
As was noted earlier, the routes
taken by females with calves during
the spring migration have been un-
known.
Twenty-three northward migrating
groups containing mothers with calves
were observed during the aerial sur-
veys (Figure 6). The earliest was
sighted 18 February, the latest on 18
May. Although the majority of those
sightings were well inshore, this may
47
be a result of the heavy sighting effort
inshore in 1972 during the times of
the northern migration. The few sight-
ings of mothers with calves late in
the season, however, were more off-
shore.
Of the 23 times mothers with calves
have been observed, in 18 mothers
and calves have been either by them-
selves or with other mothers with
calves. In only four instances were
they in the company of other adults.
This observation may be supported
in part by the fact that females with
calves are not receptive to breeding
because a female calves and breeds in
alternate years (Scammon. 1874; Gil-
more. 1961; Rice and Wolman. 1971).
Yearlings
There is still little information in
the literature on the distribution and
movements of yearling gray whales.
Hubbs (1972 pers. comm.) has ob-
served solitary yearlings migrating
south very near shore off La Jolla.
Gilmore (1960b) reported that year-
lings frequently travel with larger
adult animals on the southern migra-
tion, presumably learning the migra-
tion route, but that solitary individuals
are also seen.
Based on the growth curve of gray
whales (Rice and Wolman. 1971) and
on estimates of size range at time of
weaning (Gilmore, 1961). all whales
estimated in our surveys to be be-
tween about 6 and 9 meters (20-30
feet) long were classified as yearling
whales. The opportunity to observe
Gigi (8.26 m |27 ft] long) from the
air for nearly an hour in early March
1972 verilied the accuracy of my
previous size estimates and increased
confidence in the reliability of the
classification in subsequent sightings.
The room for error in this estimate
notwithstanding, yearling-sized whales
were observed with higher frequency
than expected (Figure 7). A total of
21 yearlings or groups of yearlings
was observed in the study area. Of
those. 16 were observed after the re-
lease of Gigi (Evans. 1972) all during
Figure 6. — Locations of
sightings of mottier-calf
groups during aerial sur-
veys. 1969-72.
Figure 7. — Locations o(
sightings ot probable year-
ling gray whales (estimat-
ed size. 20 to 30 feet) dur-
ing aerial surveys, 1969-72.
^Z"""'"'^''^i^9>
liiiiiiiiiii
'^..rrr^
"m^ 1
*--%..---— >^;;|lliil
s ;■;■;:
• NOV '^
\ ANGEL
• DEC
»~"\
:■:;:
' JAN
■\^
' FEB
• MAR
•
^
• APR
'^
\ OCEAN-
» MAY
<=^
* vicis;
Iv
*\'"m
V
- NO SIGHTS OF YEARLINGS SOUTH THIS CHART
**^i
- NO SIGHTINGS OF YEARLINGS OFFSHORE
♦ 1
- 27 AND 28 APRIL 1972 X 8 YEARLINGS FROM
A
PT CONCEPTION NORTH TO MONTERREY BAY
♦ r
34°
33°
32°
120°
the period of the northward migration.
No yearlings were seen south of the
Coronado Islands, but a total of 16
was encountered on a 2-day surve\
flight north to Monterey Bay in April.
All were within 5 km of the beach.
No yearlings were observed in the
offshore areas where other whales
were seen. Further, like cows and
calves, yearlings tended to be either
by themselves or with other yearlings.
In only 4 of the 2 I cases were year-
lings accompanying adult animals.
This absence of yearlings with adults
may be a result of the forced rejection
119°
118°
117°
h\ the mother at the lute summer
weaning in the north.
Rates of Movement
Estimates of rates of movement for
southward migrating whales have
ranged from 7.7 km/hour calculated
over the entire migration route (Pike.
1962) to 10.2 km/hour calculated
over a small segment of the route
(Cummings et al.. 1968). Rice and
Wolman (1971) used ihe times of peak
passage at two separate shore stations
to calculate the average distance trav-
48
eled In 24 hours as 185 kni. Pike
(1962) used the same calculations to
determine that northward migrants
traveled from 56-80 km/day at about
Vi-'/i the rate of southern migrants.
During this study, natural markings
on three whales observed on successive
days permitted the calculation of
speeds of movement along two areas
of the coastline. Rates of movement
of all 3 are comparable to Pike's
calculations.
Two 12-1.^ meter individuals, one
distinctly marked with white brush
markings on the tail stock and flukes,
were seen 11. 12. and 13 April 1972.
During the 49.5 hours between the
first and third sightings, they moved
approximately 129 km from the
Coronado Islands to near .San Cle-
mente. Calif., an average speed of
only 2.6 km/hour.
A 12-meter individual with a nearly
all white tail fluke and a wide white
band across the tail stock was seen
with four other animals off Point
La Jolla on 27 March 1972. The
same animal was observed again on
the 28th just northwest of Newport
Beach and on the 29th 13 km south-
west of Point Vincente. Net movement
in 44 hours was 128 km or 2.9 km/
hour.
Finally, an unusually dark yearling
observed just south of Point .San Luis
27 April 1972 had moved 64 km to
the north when it was resighted 23
hours later northwest of Point Estero.
It had moved at an average rate of
2.8 km/hour.
SUMMARY
Results of periodic aerial surveys
are comparable to those from ship
and land-based surveys in defining
the timing of migration of gray whale
populations past southern California.
Peak densities were observed in Janu-
ary for the southward and in March
for the northward migration. Over
half the population observed passed
more than 64 km offshore from San
Diego. Cows with calves were seen
from February through May primar-
ily inshore and tended to be alone or
with other cows with calves. Yearling
whales were seen inshore from Feb-
ruary through April and tended to be
solitary or with other yearlings. Fi-
nally, average speeds observed for
three individuals over small segments
of the northward migration route
were comparable to estimates based
on peak movements past shore stations.
ACKNOWLEDGMENTS
I am indebted to many people for
help with this project. John Hall and
Larry Tsunoda each flew some of the
survey flights. George Lingle and
John Moore helped summarize the
data. Raymond Gilmore. Dale W.
Rice, and William F. Perrin read the
manuscript and made useful sugges-
tions. Marita Doerflein. Marty Allen,
and Shirlee Preis typed the manu-
script.
Finally. I am indebted to the officers
and men of Carrier Antisubmarine
Air Groups 53 and 59 for their skill,
patience and helpful spirit, and to
LCDR Al Zollers for scheduling air-
craft.
This research was part of NUC IR
Project 150550 "Marine Mammal
Populations," W. E. Evans principal
investigator.
LITERATURE CITED
Adams, L. 1968. Census ol the gray whale,
1466-67. Nor.Hvalfangsl-Tid. 57:41-43.
Cumniings, W. C, P. O. Thompson, and R.
Cook. 1968. Underwater sounds of mi-
grating gray whales, EsiluiLiuiiis i;Uiiicus
(Cope). J. Acoust. Soc. Am. 44: 1278-1281.
Evans. W. E. 1972. A mobile maruie en-
vironmental survey vehicle. NUC TN No.
758.
Evans, W. E., and J. S. Lealherwood. 1972.
The use of an inslrumenled marine mam-
mal as an oceanographic survey platform.
NUC TP No. 311, 11 p.
Gilmore. R. M. 19??. The return ot the
gray whale. Sci. Am. 192(l):62-74.
. 1960a. A census of the Cali-
fornia gray whale. U.S. Fish Wildl. Serv.,
Spec. Sci. Rep. Fish. 342. 30 p.
1960b. Census and migration
of the California gray whale. (In Engl,
and Norw.] Nor. Hvalfangst-Tid. 49:
409-43 1 .
1961. The story of the gray
whale. 2nd ed. Privately published, San
Diego, 16 p.
1969. The gray whale. Oceans
1:9-20.
Gilmore, R. M., and G. Ewing. 1954.
Calving of the California grays. Pac. Dis-
covery 7(3): 13-15.
Hubhs, C. L. 1959. Natural history of the
grav whale. Proc. XVth Int. Congr. Zoo!.,
Sect. III. Pap. 46, 3 p.
Hubbs, C, L.. and L. C. Hubbs. 1967.
Gray whale censuses bv airplane in Mexi-
co. Calif. Fish Game 53:23-27.
Pike. G. C. 1962. Migration and feeding
of the grav whale iE.Mhncluins >;ibbosiis).
J. Fish Res. Board Can. 19:815-838.
Rice. D. W. 1961. Census of the Califor-
nia grav whale. 1959-60. (In Engl, and
Norw.] Nor. Hvalfangsl-Tid. 50:219-225.
. 1965. Offshore southward mi-
gralion of gray whales off southern Cali-
fornia. J. Mammal. 46:504-505.
Rice. D. W.. and A. A. Wolman. 1971.
The life history and ecology of the gray
whale {Eschruhliiis nihiisiiis). Am. Soc.
Mammal. Spec. Publ. No. 3. 142 p.
Scammon. C. M. 1874. The marine mam-
mals of (he north-western coast of North
America. John H. Carmany and Co.. San
Francisco. 3 19 p.
Wyrick, R. F. 1954. Observations on the
movements of the Pacihc gray whale.
Eschnchinis i^taicciis (Cope). J. Mammal.
35:596-598.
MFR Paper 1055. From Marine Fisheries Review, Vol.
36, No. 4, April 1974. Copies of this paper, in limited
numbers, are available from D83. Technical Information
Division, Environmental Science Information Center,
NCAA. Washington, DC 20235.
49
MFR PAPER 1056
A Note on Gray Whale Behavioral
Interactions with Other Marine Mammals
J. S. LEATHERWOOD
With the exception of reports of
killer whales, Orciniis urea, attacking
gray whales. Esihricltuus nihti.siii\.
(Scammon, 1874; Andrews. 1914;
Gilmore. 1961; Burrage. 1964; More-
John, 1968; and Baldridge. 1972) there
are no accounts in the literature on
the behavioral interactions between
gray whales and other marine mam-
mals. During aerial surveys of south-
ern California cetaceans, (Leather-
wood. 1974). I often observed gray
whales in close association with other
marine mammals (Figure I). Though
the abundance of all these species in
the area during the winter and spring
makes coincidental association likely.
the following incidents represent be-
havioral interaction;
Four days after her release, when
she was first relocated by aircraft, the
gray whale Gigi (Evans. 1974) was
swimming with a small group of Pa-
cific bottlenose dolphins Tiirsiops sp.
in the surf zone just north of the San
Clemente. Calif, pier. Though the
dolphins left the whale shortly after
the aircraft began to circle the area.
when first seen they were closely
clustered about the head of the gray
whale as if riding its pressure wave.
Since Gigi was housed during almost
her entire internment at .Sea World
with an Atlantic bottlenose dolphin.
iursiops iniiu alii.s. this association
in the wild may have been a result of
the captivity. However. I have ob-
served gray whales swimming with
bottlenose dolphins in the wild in six
other instances, in three of which the
porpoises were also riding the whales'
pressure waves. Further, bottlenose
dolphins arc common along the Baja
California portion of the gray whales"
migration route and in the breeding
lagoons and have been reported mov-
ing freely among California gray
whales (Evans and Dreher, 1962).
On 19 January 1972. three adult
gray whales were observed heading
southwest over Sixty-Mile Bank (lat.
32°()5'N. long. 1 18° lO'W). The entire
area was rich with birds and the sur-
face action of many schools of small
fishes and a large aggregation of odon-
tocetes (including over 1.000 northern
right-whale dolphins. Liwoclclphis
htircalis. approximately 500 Pacific
common dolphins. Dclphiiiiis dclphis.
approximately 500 North Pacilic
white-sided dolphins. Liiiicnoihynclius
ohUqKuicns. and at least 3 Dall por-
poises. Phocoenoidex dcilli) was pres-
ent. The whales were observed at
close range from a helicopter for nearly
45 minutes and dolphins and porpoises
were observed riding the pressure
waves of the whales for the entire
J. S. Leatherwood is with the
Naval Undersea Center Bio-
Svstems Program, San Uiego,
CA 92132.
time. All species were involved in the
interaction.
In addition, during this same period
gray whales were observed riding the
large glassy swells which moved
through the area. This behavior is
common among small dolphins (e.g..
1 iirsidps. Di'lphiiuis. La^enorhyiuhiis,
Lissodctphis) and is perhaps not sur-
prising for the gray whale in the
light of its reported surf-riding (Cald-
well and Caldwell. 1963),
In March 1971. several gray whales
were observed along the west side of
Catalina Island where an estimated
200 pilot whales. Glchiccpliald sp.,
were distributed in small groups from
Ben Wesson Point to the northwest
tip of the island. One gray whale was
turned belly up in the midst of a pod
of 12 or 15 pilot whales, and an adult
pilot whale was swimming over the
belly of the inverted gray whale. Both
were alarmed by the aircraft and
sounded on our approach.
In the other instances (Figure 1)
the animals were simply swimming
close to each other. With one excep-
tion, that of Gigi. all the observed
associations between gray whales and
other cetaceans involved adult whales.
Figure 1. — Locations ol
sightings of gray whales
associated with other ma-
rine mammals (1969-1972).
IURSIOPS SP
DEIPHINUSDELPHIS
GLOBICEPHALASP
PHOCOENOIOES DALLI
LAGENORHYNCHUS OBLIQUIOENS
LISSODEIPHISBOREALIS
- Z = ZALOPHUS CALIFORNIANUS
120
iir
118°
117
50
all greater than 30 feet in length.
Furthermore, in all cases the gray
whales have appeared to be passive
participants in the interaction.
LITERATURE CITED
Andrews. R. C 1914. Monographs of the
Pacific Cclacea. I. The California gray
whale {Rhiichuinecifs gUiiicus Cope). lis
history, habits, external anatomy, osteology
and relationship. Mem. Am. Mus. Nat.
Hist. (New Ser.) 1:227-287.
Baldridge, A. 1972. Killer whales attack
and eat a gray whale. J. Mammal. 53:898-
900.
Burrage, B. R. 1964. An observation re-
garding gray whales and killer whales.
Trans. Kansas Acad. Sci. 67:5.'^0-5? 1.
Caldwell, D. K., and M. C. Caldwell. 196.3.
Surf-riding by the California gray whale.
Bull. South. Calif. Acad. Sci. 62(2):99.
Evans, W. E. 1974. Telemetering of tem-
perature and depth data from a free rang-
ing yearling California gray whale, £.«7i-
richtnis rohiisiiis. Mar, Fish. Rev. 36(4):
52-58.
Evans. W. E., and J. J. Dreher. 1962. Ob-
servations on scouting behavior and the
associated sound production by the Pacific
bottlenosed porpoise (Tiir.siopi i;illi Dall).
Bull. South. Calif. Acad. Sci. 61:217-226.
Gilniore, R, M. 1961. The story of the gray
whale, 2nd ed. Privately published, San
Diego, 16 p.
Leatherwood, J. S. 1974. Aerial observa-
tions of migrating gray whales, Eschrich-
liiis robustus, off southern California ( 1969-
1972). Mar. Fish. Rev. 36(4):45-49.
Morejohn, G. V. 1968. A killer whale-
gray whale encounter. J. Mammal. 49:
327-328.
Scammon, C. M. 1874. The marine mam-
mals of the North-western coast of North
America. John H. Carmany and Co.,
San Francisco, 3 19 p.
MFR Paper 1056. From Marine Fisheries Review, Vol.
36. No. 4, April 1974. Copies of this paper, in limited
numbers, are available from D83. Tecfinical Information
Division. Environmental Science Information Center,
NOAA. Wasfiington. DC 20235.
MFR PAPER 1057
Aerial Observations of Gray
Whales During 1973
Radio communications with shore
observers permitted coordination of
observational efforts. Time, location,
numbers of whales, and behavior ob-
servations were noted for the sight-
ings and photographs were attempted
on occasion.
122* OO'W
I
MONTEREY OAV 1
N
/
/ ^^MONTEREY
^: "
CYPRESS -r
YANKEE PT,.f
AOBSERVATION
-A SITE
LOBOS ROCKS l
%
PT SUR -^
•^...
obsefved from ptore V
S§ A„.
oDierved from shore
SCALE 1 250.000
Figure 1. — The area off California observed for
gray whales. 15-23 January 1973.
PAUL N. SUND antd JOHN L. O'CONNOR
During their annual southward
migration California gray whales.
Eschrichliiis rohiisnis. were observed
between Monterey Bay and Point
Sur. Calif. (Figure 1) from an air-
craft during the period 15-23 January
1973. An aerial survey was initiated
in response to recommendations of
the Joint Naval Undersea Center —
National Marine Fisheries Service
(NMFS). Southwest Fisheries Center
Gray Whale Workshop (held in La
Jolla. California in August 1972). that
the accuracy of the annual NMFS
shore census taken near Yankee Point
be checked. The survey was designed
to compare shore observers' estimates
of numbers with those of aerial ob-
servers; to test the estimate that 95
percent of the gray whales migrating
by Yankee Point pass within 1.9 km
(1.2 miles) of the shore (Rice and
Wolinan. 1971); and to provide ob-
servations of gray whale behavior and
associations with other marine mam-
mal species. The utility of aerial sur-
veys in cetacean research has been
demonstrated by Levenson (1968) and
Leatherwood ( 1974a. b). This paper
reports on simultaneous shore and
aircraft observations and discusses
the problems inherent in each method.
METHODS
Five flights, totaling 13.6 hours,
were made between Monterey Bay
and Point Sur. Calif. (Figure I) in a
Cessna 172 flown by a professional
spotter-pilot at altitudes ranging from
150 m (500 ft) to 900 m (3,000 ft).
RESULTS AND CONCLUSIONS
From the aerial observations made
in the sector scanned by shore obser-
vers, the following points were deter-
mined: Of 24 paired observations
(individuals or groups observed by
both air and ground personnel), ini-
tial visual contact was made by a
ground observer in eight instances and
by an airborne observer in ten in-
stances. Hence, ground and aircraft
observers apparently were equally
adept at initially sighting whales. Of
the 24 paired sightings, the aerial
observers were able to correct the
Paul N. Sund is with the Pacific
Environmental Group, National
Marine Fisheries Service,
NOAA, Monterey, CA 93940.
John L. O'Connor, P.O. Box
1942, Newport Beach, C A 92660.
51
numbers recorded by the shore ob-
servers six times. In three instances of
poor (white caps and 4-6 foot swells)
sea state conditions, on the other hand,
the aerial observers were unable to
contirm groups or individuals sighted
by the shore observers. These data
suggest that, although aerial observa-
tions may be more directly limited by
sea conditions, they are useful in
quantifying the number of whales in
groups. Further, resolution of num-
bers present is faster from the air
than from shore. (It often takes the
shore observers up to 30 minutes to
determine their count for a given
group — during which time the in-
dividuals in the group may dissociate
or join with others.)
Resolution of numbers of whales
in groups is more rapid and apparent-
ly more accurate from the air than
from shore. With a professional spot-
ter pilot working a limited area —
such as that scanned by the shore ob-
servers— in good sea state conditions,
essentially no whales will pass unno-
ticed. "Misses" by the aerial observers
were due to leaving the area premature-
ly in order to accomplish other tasks;
had the aircraft been consistently in
the shore observers' area (and immedi-
ately outside to prevent unnoticed
passage of individuals offshore) none
would have gone unrecorded.
The aerial observers made ."^O ob-
servations of whales involving 149
animals. All these observations oc-
curred within 7 miles of the shoreline,
even though the area surveyed ex-
tended to 25 miles seaward. Of these
sightings. 98 percent were within 5
miles of shore. 96 percent within .^
miles, and 94 percent within I mile.
Distances were estimated by making
timed runs at constant speed from
pi>sitions offshore to the coastline.
The observations of this study tend
to confirm Rice and Wolmans state-
ment that 95 percent of the whales
pass within 1.9 km (1.2 miles) ot the
shore near the Yankee Point site.
Gray whales have been reported
interacting with other marine mam-
mals by Leatherwood (1974b). but
during this study no other marine
mammals were observed "associating"
directly with gray whales. Feeding
behavior was observed on two occa-
sions. A calf was seen accompanied by
an adult. These two latter observa-
tions are of particular note and the
senior author intends to publish the
details elsewhere.'
' Sund. P N Manuscript Evidence of feeding
during migration and of an early birtti of the
California gray whale
LITERATURE CITED
Leatherwood. J. S. l'J74a. Aerial observa-
tions of migrating gray whales. Eschrich-
liiis rubusitts, otT southern California
(1969-1972). Mar. Fish, Rev. 36(4):45-49.
. 1974h. A note on gray whale
behavioral interactions with other marine
mammals. Mar. Fish. Rev. 36(4):49-?0.
Levenson, C. 1968. Factors aftecling bio-
logical observations from the ASWEPS
aircraft. U.S. Naval Oceanogr. Off., In-
formal Rep.No. 68-102, 6 p.
Rice. D. W., and A. A. Wolnian. 1971.
The life history and ecology of the gray
whale {Eschnchtnts nihusius). Am. Soc.
Mammal., Spec. Publ. No. .3. 142 p.
MFR Paper 1057. From Marine Fisheries Review, Vol.
36. No. 4. April 1974. Copies of this paper, in limited
numbers, are available from D83. Technical Information
Division. Environmental Science Information Center,
NOAA, Washington. DC 20235.
MFR PAPER 1058
Telemetering of Temperature and Depth Data
From a Free Ranging Yearling California Gray
Whale, Eschrichtius robustus
W. E. EVANS
ABSTRACT
//; /y6(S' //((■ iiinlinr iiiiluilcti a .scries nf Miii.lics iisiiii; icidic iniii.sniillcr.s lo
jollow ilw nidViincnls iiiitl sliidy ilw cliviiii^ hchiivior ol snuill Icollwil whales.
This paper dc.si ril'cs the miulifu iilit'irs tij this ccpiipniciU necessary lo use this
Icchniquc on larj^cr wTnilcs. in this case a yciirliiii; California ,t,'/((y wliale,
Eschrichtius robustus. In addition lo the iransinission of positional data, i.e.
azinuilh anil deplh of dive, the ittsininicnlalion pnekaf^e used in this study was
desii;ncd lo trausnui environmental data (tcinperaturc-al-depihl. The animal
used in this study, a Jemalc E. robustus, U(/.v captured on /.■? March 1971. in
Smnimon's Lagoon. Baja California Siir. Mexico, hy Sea World, Inc., San Diei.;o,
and released on 13 March 1972. at hit. 32''4I.5'N. lonn- 1 17^^ 20.5' W {off Point
Loina. San Die.i^o. Calif.) hy the Naval Undersea Center (NUCI. San Diei;o.
Rtulio contact was maintained with the animal iinlil .'^ May 1972. Deplh oj dive
and leinperalure-al-deptli data w ere idnlniuouslv mniiiloreil for a 24-hoiir period.
INTRODUCTION
The present stud\ is an extension
of a 6-year research program designed
io evaluate the feasibility of usmg
medium-si/ed lo large cetaceans, in-
strumented with a combination data
collection and transmission system.
to measure physical oceanographic
parameters at various depths, and to
evaluate the relationship of these
parameters to cetaceans' movement
patterns and secondary productivity
(Evans. 1970. 1971. in press).
Because of the impending release
of a yearling California gra\ whale
52
W. E. Evans is with the Naval
LJndersca Center. San Diego,
C A 92132.
(Gigi II) captured by Sea World, Inc.,
San Diego, on 13 March 1971, in
Scammon's Lagoon. Baja California.
Mexico, and the timing of the northern
migration of CaMfornia gray whales
(March-Aprill, the program was ac-
celerated to take advantage of this
opportunity. The prototype data trans-
mission/acquisition system had been
designed and bench-tested in anticipa-
tion of tests on a Pacific pilot whale.
ClohiccphaUi cf. sctiniiiuini. in mid-
summer 1972. The unit was repack-
aged and the test dates subsequently
moved up to coincide with the planned
release of the Sea World captive year-
ling gray whale which was scheduled
for 13 March 1972. It was then field
tested attached to the ,Sea World gray
whale when she was relased at 0905
hours at lat. 32°4I.5'N. long. 117°
20,5'W into a group of four to five
California gray whales moving north.
DATA PARAMETERS
Since one of our primary purposes
for using a data system attached to a
cetacean was to measure environ-
mental parameters associated with the
animafs movements below the air-sea
interface, the instrumentation used
must indicate the depth at which the
measurement was made. The follow-
ing parameters were considered as po-
tential indicators of productivity and
important correlates of cetacean
movement;
1. Temperature at depth.
2. Ocean current speed at surface
and at depth.
3. Salinity-derived from conduc-
tivity measurements
4. Dissolved gases:
a. O2:
b. No;
c. FreeC02.
5. Light;
a. Absorption loss due to molec-
ular absorption, particulate
matter;
Figure 1. — Block diagram of telemetry transmitter attactied to yearling California gray wtiale (Gigi).
b. Backscattering from particu-
late matter;
c. Light level at depth.
After consideration of all these
parameters, temperature was selected
as the most desirable for this phase of
the program because; 1) methods of
measurement are straightforward elec-
tronically. 2) considerable bathyther-
mal data exist for the California C ur-
rent region, 3) data transmitted from
the instrumented animal could be
easily checked by use of currently
available expendable bathythermo-
graphs, and 4) a great deal of data
relating the thermal structure of the
water columns to primary and secon-
dary productivity are available in the
scientific literature (e,g. Eckman,
1953).
INSTRUMENTATION
Data Transmission System
The data acquisition system mount-
ed on the yearling gray whale pro-
vided measurement of the depth of
each dive and the water temperature
at that depth, and served as a radio
beacon for tracking. Data measured
was telemetered by an 1 I meter trans-
mitter (27.585 megaHertz) to either a
surface vessel, shore station, or air-
craft-based receiving set which would
also demodulate the data being trans-
mitted. Directional information for
tracking was obtained by a special fast
response automatic direction finder
developed several years ago specifical-
ly for this type of application (Ocean
Applied Research Corporation, San
Diego, Model ADF210).i
A block diagram of the telemetry
transmitter is shown in Figure I . Func-
tion and operation are as follows;
Pressure is measured by a semi-
conductor strain-gauge bridge excited
with constant current. The output
voltage is amplified by three opera-
tional amplifiers connected in an
"instrumentation amplifier" configura-
tion and the peak pressure reading
stored (remembered) on a capacitor
which is followed by an insulated-gate
field-effect transistor (LET). This
peak detector is inside the feedback
loop of the amplifier which maintains
accuracy and also yields a digital
zero at point A when pressure is de-
creasing from the peak depth. This
level is used to hold the temperature
reading.
' Use of trade names in this publication does
not imply endorsement of commercial products
by the National Marine Fisheries Service
53
Temperature is measured by a therm-
istor composite which is pressure
protected in a thin-wall stainless steel
tube. The thermistors' conductance is
measured by an "operational trans-
conductance amplifier" (OTA) whose
output is gated by the digital signal
from the pressure sensor. The output
of the OTA drives a capacitor which
serves as a temperature-reading mem-
ory. It is also followed by an insulated-
gate FET whose high input impedance
prevents memory discharge.
These two voltage analogs are con-
verted to frequency analogs by voltage
controlled oscillators (VCO). The fil-
tered outputs of the VCO's are
summed and the resulting composite
fed to the amplitude modulator of the
3-watt peak transmitter.
A programmer is also included to
provide a 4-second data transmission
time when the animal first surfaces
followed by a series of short pulses
which are adequate for the tracking
system. A seawater connection be-
tween the antenna tip and the instru-
ment case generates a delayed reset
for all capacitor memories and the
programmer.
In the package used on the yearling
gray whale two batteries were included
Figure 2. — Photograph of the Ocean Applied Re-
search Corporation data transmitter Model WDT-
920 attached to the Sea World yearling gray
whale (Gigi) just prior to release. (Photo courtesy
of J. S. Leatherwood.)
Figure 3. — Aerial photograph of lest animal taken on 16 March 1972. as she was swimming through
kelp beds off San Clemente, California. (Note kelp trailing behind the transmitter package.) (Photo
courtesy of J. S. Leatherwood.)
in the system. One had a capacity of
1.^ ampere hours and was used to
power all electronics which were on
when the animal was at the surface.
The second, smaller, battery, which
had a 1.2 ampere hours capacity,
powered the depth and temperature
instrumentation continuously. The ex-
pected life of the smaller battery was
approximately I month while the
larger battery with its greater capacity
and reduced duty cycle should con-
tinue to provide tracking transmis-
sions for as much as 9 months. The
entire system packaged and attached
to the whale is shown in Figure 2.
During the first month of operation.
V
'/, WAVE
MARINE ANTENNA
BAND
PASS
FILTERS
TEMPERATURE
DISCRIMINATOR
n
DEPTH
DISCRIMINATOR
_
-
DIGITAL
READ OUT
TAPE
RECORDER
Figure 4. — Block diagram of the telemetry data
receiving and recording system.
performance of the Instrument/Bea-
con package was satisfactory with the
notable exception of transmission
range which was initially more than
25 miles. Subsequent tests indicate
that the antenna, which is a top loaded
stainless steel whip antenna, had sus-
tained some damage. The animal was.
on two occasions, observed swimming
through kelp and kelp was seen hang-
ing on the antenna. Figure 3. Cali-
fornia gray whales are also known to
rub on the bottom, a behavior which
could have abraded or even severed
the loading coil from the antenna,
drastically reducing its radiation ef-
ficiency. The estimated useful range
of the damaged system was on the
order of 10 miles.
The original antenna design em-
ployed on beacon transmitters for ma-
rine mammals (specifically porpoises)
was an adaptation from a design which
had been in use for some time on
radio beacons used for the recovery
of oceanographic instruments. It con-
54
Figure 5. — Automatic direction finding antenna (loops) and data acquisition antenna (whip) attached
to the belly of a U.S. Navy S-2 tracker aircraft. (Photo courtesy of J. S. Leatherwood.)
sisted of a solid fiberglass tapered rod
onto which was wound a conductor
and loading coil. An aluminum tip
served both as a seawater contact and
as a section whose length could be
trimmed tor peak field strength. A
proprietars coating protected the con-
ductor and coil from seawater. The
antenna was entirely successful on
Delphiinis species (Evans, 1971). but
problems were encountered when a
similar design was used on captive
whales such as pilot whales and killer
whales. The captive whales invariably
broke the antennas by rubbing on
structures or boats and in the instance
of the release of a pilot whale into the
wild, the antenna was broken by sea-
weed. Subsequently, a spring-wire
antenna was designed which could be
severely bent without catastrophic
damage and has been used success-
fully on the aforementioned whales
(Martin. Evans, and Bowers. 1971).
This type of antenna was used on the
gray whale pack. Subsequent simula-
tions of various types of damage to
this antenna indicate that modifica-
tions would be in order before em-
ploying this type of antenna again.
Specifically, the arrangement of the
spring at the base should be changed
to allow the antenna to be bent double
against the transmitter case without
damage. The antenna should be length-
ened somewhat to reduce the variation
in impedance for a given variation in
the relative position of the ground
plane (sea surface) and the loading
coil should be fully encapsulated in
the nonmetallic antenna's structure to
completely eliminate the abrasion
damage potential.
Data Receiving System
The data receiving and recording
system illustrated in Figure 4 was
originally tested on board the NUC
RV Cape. Subsequent to the initial
tracking and data acquisition attempts
following the release of the whale on
13 March 1972. the system was placed
on board a U.S. Navy S-2 tracker
aircraft. The antenna mounting con-
figuration used on this type of aircraft
is shown in Figure 5. The whip anten-
na is shown in the retracted mode. The
loop antenna used in conjunction with
the automatic direction finding system
is adjustable and was aligned prior to
every flight by using a shore-based
radio beacon.
Several relocations of the animal
were made using this system. The short
transmission range of the damaged
transmitter system attached to the
whale seriously limited the acquisition
of temperature and depth data from
the aircraft-mounted system. Tests
>-*%► .„
r*-^?25B»-^
/-""':.' pz::^
f^%
-P
Figure 6. — California gray whale breaking the
the back. The normal sequence of a blow is
note exposure of the dorsal ridge. (Photo courtesy
surface exposing only the head and fore-part of
shown right to left at the bottom of the figure,
of J. S. Leatherwood.)
55
Gray Whale Relocation Records
Enlargement
Of ThiB Area
Figure 7. — Map of California coastline showing
(Gigl) 13 March-5 May 1972.
conducted following the release of the
whale using a similar data transmitter
with one-third the power of the system
used on the test animal and a modified
antenna have yielded data acquisition
ranges up to 40 nautical miles.
RESULTS
During the initial 2 hours after re-
lease of the test animal, signals were
very intermittent and seldom longer
than 2 seconds in duration. Ohserva-
tions lead us to believe this was a
behavioral problem since the animal
frequently broke the surface of the
water showing onl\ her blow hole and
mid-portion of her back as illustrated
in Figure 6. Since the data transmitter
was mounted on the dorsal ridge (on
the last half of the body) the antenna
either did not break the surface of the
water, thus no transmission, or only
the tip of the antenna broke the sur-
face, resulting in a very short duration
transmission.
locations o( the Sea World yearling gray whale
This resulted in limited data recep-
tion during the first 2 hours after re-
lease and subsequent loss of the ani-
mal's location and movement pattern.
Those signals over 2 seconds in dura-
tion that were received during this
time period did indicate temperature-
at-depth values reasonable for the lo-
cation and time of year (e.g.. above
20 meters temperatures of 13°-I4°C
and below 20 meters a temperature
of 7.4C'). A 20-meter isothermal layer
is not uncommon at this location.
Since the quality of radio signal ac-
quisition was quite poor, the search
from the RV Cupc was abandoned in
favor of an aerial search. The animal
was relocated on i.^i March between
1300 and 1500 hours on a bearing of
320°T south of Oceanside. The animal
was relocated again on 16 March
close inshore off San C'lemente. Calif.,
working slowly north. The photograph
shown in Figure 6 was taken at this
time. On this flight and those that
followed, although the animal could
be easily located, acquisition of use-
able temperature-at-depth data was
limited I) by the long time interval
between adequate exposure of the an-
tenna, and 2) by the apparent short
range of the transmissions received.
Areas of visual relocation and radio
contact from 16 March 1972 to 5 May
1972. are illustrated in Figure 7.
After a period of 5 to 6 days, the
animal's swimming pattern changed
and longer and more frequent trans-
missions were being received. In order
to verify these observations and. if
possible, to collect temperature-at-
depth data over a 24-hour period, the
RV Ciipc left San Diego at approxi-
mately 1600 hours on 20 March 1972
for the Dana Point-San Clemente,
Calif, area. At IS4() hours. 3.'^ nautical
miles from San Clemente. Calif., we
acquired weak signals from the animal
bearing 340°T. At 2300 hours, signal
level had increased and we were re-
ceiving bearing and temperature-at-
depth data. Initial data indicated tem-
peratures of 12°-I4°C at depths of
l.'^-20 meters. At 2350 hours the
animals diving behavior changed and
indicated some dives to depths of 170
meters. Triangulation placed the ani-
mal at a location approximately on the
100 fathom curve. 1.7 nautical miles
off Laguna Beach. Calif. (Aliso Can-
yon). Although the depths recorded
at this location were realistic if the
animal was diving to the bottom, the
water temperatures at those depths
appeared to be anomalous. At this
HktlMUM DOWH TIME - 16 m
Figure 8. — Mean down times and depths of dive
as a function ol time of day recorded 20-21
IVIarch 1972.
56
time of the year one would expect sur-
face temperatures between 12° and
i4°C. and temperatures at 170 meters
of 7°-8°C or less. Data from the
animal, however, indicated tempera-
tures-at-depths of 100-170 meters
ranging from 10° to 14°C. If these
levels were indeed accurate, a signifi-
cant temperature inversion layer was
present. This cruise was terminated at
2100 hours on 21 March 1972. after
having recorded data for approximate-
ly 24 hours. In addition to the temper-
ature-at-depth data, the following
observations were made:
1. The animal was offshore 1-2
nautical miles after sunset, moved
inshore 100-200 meters from the
beach post -sunrise.
2. The diving pattern at night was
regular, as compared to an erratic
pattern during daylight hours.
3. The animal was observed in
Dana Cove along with three other
gray whales of a similar size range.
All four animals left the Cove in late
afternoon.
4. The mean time between trans-
missions was significantly longer from
1600 to 0400 hours than during the
remainder of the day (Figure 8).
Since transmissions were more pre-
dictable after sunset, our aircraft data
acquisition flights were scheduled at
night. Under this plan, data were col-
lected on 28 and 29 March from the
S-2 tracker aircraft in the vicinity of
Dana Point, Calif. The recorded data
from these flights indicated dives of
50-80 meters and temperatures of
I2°-14°C. The observations that the
animal moved offshore (1-4 miles) at
night were verified.
No readily apparent explanation
was available for the relatively high
temperatures recorded at depth on
20-21 March. To check on these
measurements, the RV Sen Sec was
sent to the Dana Point area to work
from 6 April 1972 through 10 April
1972. equipped with an expendable
bathythermograph (XBT) system ca-
pable of measuring temperature versus
depth over a range of 0°C-30°C to
depths of 450 meters. During the
Figure 9. — A composite of three expendable
bathytliermograph plots collected 2 miles oil
Dana Point, Calilornia, at 1700 hours on 10
April 1972.
period 6-7 April, the area south of
Dana Point was searched and no con-
tact was made with the animal, al-
though one small 7-8 meter whale was
sighted. The XBT data, however, in-
dicated surface temperatures of approx-
imately 13°C which dropped to 6°C
at 300 meters, with no obvious Iher-
mocline or temperature inversion.
Late on 7 April, a search was made
north of the Dana Point area by auto-
mobile and signal acquisition was
made from the Huntington Beach pier
at 1600 hours. Observers on the pier
claimed to have sighted a small Cali-
fornia gray whale with a radio pack
swimming north 100 meters off the
end of the oi'^'- at 1000 hours on that
same day. Our signal acquisition was
on a bearing of 280°T which would
1 1
J
-
••-"(•
-
-
J
lOCAIION. \.J mlieS OFF
LAGUNA lEACH.
'
j
CAlIf
Il*tl. IBOS
/
■ ANGE TEMPCRATUIt
ATDCPTM RCCOIDfD
/
HIS, 10 m 7] TO
0830, 31 in 73
Figure 10. — A composite of three expendable
bathythermograph plots collected 1.7 miles off
Laguna Beach, California, at 1800 hours on 10
April 1972, compared to temperature versus depth
data collected from Gigi on 20 March 1972.
place the animal between Seal Beach
and Santa Catalina Island.
On 10 April 1972. the RV Sea See
started a search north of Dana Point
at 1600 hours. The first XBT station
was at 1700 hours. 2 miles off Dana
Point. A plot of temperature versus
depth representative of the three mea-
surements taken at this location is
presented in Figure 9. The vessel
moved north to approximately the
same location where the temperature-
at-depth data were collected from the
whale on 20 March. Of major interest
here is indication from the XBT data
that a rather significant temperature
inversion did exist in this area. A plot
of the XBT station taken at 1800
hours on 10 April. 2 miles off Laguna
Beach compared to the temperature-
at-depth data transmitted from the
whale is illustrated in Figure 10. It
should be noted that although the re-
lationship between the XBT and whale
data appears to be comparable at
depths of 100-200 meters, these are
preliminary data. Conclusion should
not be made without further verifica-
tion of the thermal inversion shown
in Figure 10. It is possible that the
outfall of the San Onofre nuclear
power plant north of this location is
dumping warm water back into the
sea. It has also been shown that at
this time of year the mean geostrophic
flow at 200 meters has northbound
components (Wyllie, 1966). Depend-
ing on the temperature, volume, and
depth of the San Onofre effluent it is
not inconceivable that inversions such
as that shown in Figure 9 could result.
This, however, is pure speculation at
this point.
ACKNOWLEDGMENTS
This study was accomplished under
funding from NASA. Defense Pur-
chase request A-70496A, RTOP 160-
75-81, Task 01, Ames Research Cen-
ter, Moffett Field, California. Addi-
tional funding support was received
from the Naval Undersea Center, San
Diego, California, Independent Re-
search project ROOO-OI. All of the
electronic equipment used to instru-
57
nient the whale and subsequently
track and obtain data Ironi it was de-
signed and manufactured by Ocean
Applied Research Corporation. San
Diego, California. Hugh Martin and
Romaine Maiefskl. both from this
organization, actively participated in
the attachment of the instrumentation
to the animal and the initial stages of
tracking. J. S. Leatherwood. J. Hall.
Bruce Parks, and L. McKinley. of the
Naval Undersea Center. .San Diego.
California, and the Commanding Of-
ficer of the RV Cape and his crew
were directly instrumental in the suc-
cess of this project. The radio contact
with the instrumented whale on 5 May
1972 was accomplished by Paul Se-
besta. NASA Ames Research Center.
Moffett Field. Calif., using equipment
supplied b\ the author.
due to the destruction or displacement
of melanin in the epidermis of the
area treated.
LITERATURE CITED
Eckman. S. 1953. Zoogeography ol llie
sea. Sidgwick and Jackson Ltd.. Lond..
417p.
Evans. W. E. 1970. Uses of advanced
space technology and upgrading ihe future
study of oceanology. AIAA 7th Annual
Mtg. and Tech. Display. Houston, Tex.,
Pap. No. 70- 1:?.^, p. I-.^.
. 1971. Orientation behavior of
delphinids; Radio telemelric studies. Ann.
N.Y. Acad.Sci. 188:142-160.
. In press. Radio lelenietnc
studies of two species of small odonlocete
cetaceans. In W. E. Schevill (editor).
The Whale Problem. Harvard Press,
Cambridge, pp. 385-394.
Martin, H., W. E. Evans, and C. A. Bowers.
1971. Methods for radio tracking marine
mammals in the open sea. IEEE Eng. in
the Ocean Environ. Conf.. p. 44-49.
Wylhe. J. G. 1966. Geostrophic How of
the California Current at the surface and
at 200 meters. Calif. Coop. Oceanic Fish.
Invest. Atlas No. 4. ,\iii + 288 charts.
MFR Paper 1058. From Marine Fisheries Review, Vol.
36, No. 4, April 1974. Copies of ttiis paper, in limited
numbers, are available from D83. Technical Information
Division, Environmental Science Information Center,
NCAA. Washington. DC 20235.
MFR PAPER 1059
POSTSCRIPT
During the period 2 January 197.^-
21 March 1973. the author investigated
37 reported resightings of Gigi. Al-
though most of these reports did not
check out, on 5-6 January, a Captain
Paul Roth. USN. and a Mr. and Mrs.
Sherwood of San Diego independently
described behavior of a 9-10 meter
California gray whale sighted inside
the kelp off the Sunset Cliffs area of
Point Loma. San Diego. California.
In both cases the whale, light in color.
approached close to small vessels less
than 10 meters, rolled, and frolicked
around. On 15 March we received a
report from the MV Loiii; Bench
Prince that a whale of similar size
and with white tail Hukes (see Figure
3) and a fiO cm X 60 cm square white
scar behind the blow hole was sighted
frolicking around the vessel by 178
whale watchers. The location of this
sighting was 3-4 miles off Point
Fermin. This latter sighting is es-
pecially interesting since on 6 March
1972, one week prior to release, Gigi
II was branded using cryogenics with
a 60 cm x 60 cm mark, midline on the
back just posterior to the blow hole.
This form of marking, called "freeze
branding. " results in a white scarring
Capture and Harnessing of Young
California Gray Whales, Eschrichtius robustus
KENNETH S. NORRIS antd ROGER L, GENTRY
ABSTRACT
1 lii\ paper reptirls mi the tleuiils i>f capture , liarnessiiii;. ircukiiii;. ami lianiess
release for three suckluiii i:ra\ u/ki/cv. These lesls are ihe firsl steps in a pr(ii;raiii
Id ilevelop new means af clala acqiiisilion and recovery jri'in wliales diirini; iheir
mii^raiiinis. I) is hoped hy tliese means u> develop new infornuilion ahoiil
popiilaliini routes and hence population immhcrs to assist nianai;einent. Capture
was hy tail noosiiii^ and liead nelliiii; from a fishins; vessel equipped with a
swonljisli plank. The liarness. placed on the inptive ashore, was held in place
over tlie pectoral fins and hcuk hv means of a pair of inelal plates held together
hy a \oliihle nnc.;ne\inm holt . Trackiiii^ was hy radio.
INTRODUCTION
Informed whale management re-
quires adequate knowledge of popula-
tion numbers. Uncertainly about mi-
gratory pathways and population mi.\-
ing makes determination of such num-
bers uncertain for some whales such
as the humpback {Mci^aptera novaeaii-
Siliae). the blue whale {Balaenoptera
iniisciihis). the (in whale iBalaenop-
lerii phxsaliis). and the minkc whale
{Balaenopiera aciorostraia). Thus pre-
cise information on migration routes
of these and other marine mammals
would materially assist in the develop-
ment of sound management practice
(Anonymous, in press).
In spite of decades of work with
Discovers and other tagging methods
(Clarke, 1957) our knowledge of whale
migration remains highly incomplete.
Because such information is needed
for some protected species, new tag-
58
Figure 1. — Map ol capture locality. Nuinbers
indicate the capture sites for the ttiree harnessed
animals.
ging methods that do not require kill-
ing are now required. These methods
seem to fall into two categories; (1)
those involving the capture of whales,
placement of harnesses and equipment
on them, tracking along the whale's
route, and subsequent release and re-
covery of data packages; and (2) those
involving placement of data or tele-
metering packages on whales without
capture, followed by tracking.
The first method will allow data
collection from a few animals, while
the latter will presumably allow less
complete data collection from more
animals and from those species that
cannot be captured. The tests de-
scribed here are of the first sort; that
is. they involve capture and harness-
ing. The experiments of Evans (this
number of Marine Fisheries Review)
with Gigi are also of this sort, though
surgical attachment rather than har-
nessing was used.
We chose our subject, the California
gray whale, because large numbers
of suckling calves are available in
their Mexican breeding lagoon during
January and February of each year
and because the calm working condi-
tions in the lagoon would assist these
preliminary tests. We expect that the
majority of results obtained on this
relatively well-known animal will be
applicable to more oceanic species.
Our tests were restricted to capture,
harnessing, and very short term track-
ing, since we expected that our results
would require harness redesign prior
to long-term tracking. This proved to
be the case.
We attempted to capture suckling
animals only because of the obvious
dangers and seamanship problems pre-
sented by adult whales. The rationale
supporting this choice is that a suck-
ling calf, harnessed and instrumented,
should keep station with its mother
and. hence, give a true migratory
route.
To our knowledge five baby gray
whale captures or handlings have
been reported. Eberhardt and Norris
( 1964) report working with a stranded
baby gray whale in Scammon's Lagoon.
Robert Eisner (pers. comm.) detailed
a capture of a baby gray whale in
Scammon's Lagoon from a small cata-
maran by use of a superficial harpoon
followed by netting. David Kenney
(pers. comm.) directed the capture of
Eisner's animal and the capture and
transportation of Gigi. the gray whale
calf caught in Scammon's Lagoon and
held for 12 months in Sea World.
The latter whale was captured with a
tail noose from a small fishing vessel
equipped with a bow plank. The ship
was reportedly damaged slightly by
the mother when the baby was brought
alongside. Theodore Walker (Cous-
teau. 1972) is shown manipulating a
stranded baby gray whale in circum-
stances much like those discussed by
Eberhardt and Norris (1964). Spencer
(1973) reported on the drug-assisted
capture of adult whales in Scammon's
Lagoon.
THE STUDY SITE
We chose northern Magdalena
Bay. Baja California Sur. Mexico,
near Boca de Soledad for our work
because of an abundance of whales
living in a system of shallow bays and
rather narrow channels and because
Kenneth S. Norris and Roger L.
Gentry are associated with the
Coastal Marine Laboratory, Uni-
versity of California, Santa
Cruz,' Santa Cruz, CA 95064.
the Mexican government has recently
declared the better known Scammon's
Lagoon (Laguna Ojo de Liebre) a
whale reserve. Headquarters were es-
tablished in the small government
cannery town of Lopez Mateos. which
fronts on the main lagoon channel 8
km southeast of Boca de Soledad
(Figure 1).
In this region the channel is about
800 m wide and averages 1 1 m deep
in mid-channel. To the west a low
ridge of dunes separates the lagoon
from the sea. The shore along the
dunes drops precipitously into deep
water. The eastern bank is typically
bordered with dense mangrove thick-
ets often cut by shallow bays and
channels. The shore along the man-
grove coast usually shelves gradually
over a broad tidal flat to the main
channel. This difference in bottom con-
tour proved crucial to capture and
harnessing.
While whales were found through-
out the deeper parts of the channel,
one concentration occurred just in-
side Boca de Soledad and another oc-
curred at a broad expanse of water just
north of Colina Coyote (see Figure 1).
It was here, or somewhat closer to
Lopez Mateos. that our captures took
place. Our counts showed approximate-
ly 86 whales in residence in the entire
channel system. Most were mothers
and young, but a few males were pres-
ent, as indicated by copulations ob-
served inside the channel.
WHALE CAPTURE
AND HARNESSING
Capture of suckling gray whales
proved to be rather simple, once the
basic techniques were established. Four
whales were netted in 3 days (27-29
January 1973). One was released be-
cause it was clearly too large for our
59
harnesses. The other three were suc-
cessfully harnessed, released to their
mothers, and tracked. Capture was
performed from the swordfish boat
Loiison. a 15-m vessel equipped with
a II .5 m welded aluminum pipe pulpit
projecting from its bow. During cap-
ture Captain Tim Houshar occupied
the basket at the end of the pulpit,
while the helmsman steered from a
remote station atop the crow's nest.
The vessel was maneuvered behind a
whale pair, attempting to place the
netman in the pulpit over the animals
as they surfaced to breathe. At the
same time another crewman in a
speedboat zigzagged around and in
front of the animals in an attempt to
direct and distract them. This attempt
succeeded often enough that surfacing
whales rather regularly allowed the
pulpit to pass over them. The tendency
to surface beneath the pulpit varied
rather widely from pair to pair and
seemed most consistent in mothers
with small young.
Once a pair surfaced under the pul-
pit a noose of 1.25 cm nylon line was
placed over the small animal's head by
means of a large metal hoop cut
through at its outer margin and held
together inside a piece of plastic
tubing (Figure 2). The rather slow
speed of the whales (usually less than
7 knots) and the relatively long time
they spend at the surface during res-
piration make this a reasonably sim-
ple process.
At this point the nylon noose which
was tied to the metal hoop with light
twine was pulled loose. The hoop
separated over the animal and was
pulled away, leaving the noose to slip
back to the tail stock of the little whale.
Another crewman on the pulpit pulled
the noose tight over the tail stock. The
noosed young took out a modest
amount of line, usually less than 100
m. before the line was belayed around
a Samson post. The young did not
dive for extended periods (less than
1 minute) but towed the vessel for a
time in this position. The mother
always stayed in close attendance.
Figure 2. — Capture of the baby whale. Note hoop and noose being placed over the baby. Note also
the swordfish plank which is maneuvered over the mother-young pair.
often sliding over the line or coming
up underneath it. At times she lifted
the young on her snout or back, and
occasionally she thrashed at the re-
straining line with her flukes.
Once the young animal began to
slow somewhat, it was brought back
under the pulpit by bringing in line.
The mother came with it and swam
under the pulpit or slightly off to the
side. Never did a female attempt to hit
the boat or the pulpit, though our
small sample may not be representative.
A head net bag of 5 cm nylon mesh,
also containing a noose of 1.25 cm
nylon line and similarly positioned
on a hoop frame, was placed over
the baby's head. Optimally this net
was deep enough to extend from the
tip of the snout to just posterior to the
pectoral flippers. In practice our nets
were too small for all but one animal
and placed the noose anterior to the
pectorals. Even so. the noose did not
slip loose.
With lines fore and aft the young
animal was severely hampered and
could be pulled in rather easily by
hand. During this time the boat and
skiff had been maneuvering the pair
toward the east bank and its shallow
shelf.
Two plastic trash barrels containing
the coiled head and tail lines were lift-
ed into a waiting skiff and payed out
to the restrained whale until the shallow
shelf was reached. Then the lines were
taken ashore and the men. usually
four to six. pulled the baby sideways
onto the shelf. Usually the mother's
efforts were strenuous at this time, and
occasionally she looped the line over
her body or tail giving an irresistable
pull, but always she rapidly slipped
free and the baby could be towed in
again. The baby was beached in about
0.7 m of water. 10 m or so from the
shelf edge. The mother was unable to
enter such shallow water, though she
did patrol the shelf edge, and in one
case partially stranded herself, seem-
ingly in an attempt to reach the baby.
Thus protected from the obvious ire
of the mother, it was rather simple to
60
place ihc harness on the baby. The
danger from the mother was made
clear when a crewman began working
within a few meters of one. The female
whale lifted her tail, bent it back and
thrashed the flukes around in a semi-
circle, horizontal to the water surface.
She missed the man by quite a dis-
tance but the force of the blow was
enough to send a sheet of water over
everyone nearby.
The baby remained rather quiet
during the harnessing process. The
harness was usually slipped on under
the snout and v/orked posteriorly to
the pectorals which were then inserted
through the harness. The harness was
then tightened in place until snug over
the baby's body. At this point, timing
for harness release began as a corrosi-
ble magnesium bolt which held the
release mechanism began to corrode
away in salt water.
Three or four men pushed the baby
back into deep water over the shelf
taking care to avoid the mother. In all
but one case she was nearby and
quickls took up station with her off-
spring. In one case the mother left
before the baby was launched and
was a kilometer or so down the bay
shore when the baby began to swim
in deep water. This baby cruised
quietly for a short time and then, when
about 300 m from the mother, turned
as if on a signal and raced toward her.
The mother did the same, turning
toward the baby and beginning to
swim rapidly. Once they were near
the mother circled the baby, thrashing
the water with her flukes. It was prob-
able that an acoustic recognition sig-
nal was involved. This young animal
had been emitting short low frequency
signals while stranded. Even if the
young did become separated from the
mother by some distance, chances for
reunion remained excellent because
of the restricted channels available
for swimming.
In all cases the presence of the har-
ness had no visible effect on the be-
havior of the mother-young pair.
HARNESS DESIGN
The harness was constructed of
four lasers of one-way stretch Lino
241,' commonly used in fabricating
girdles and corsets, that permitted
expansion and contraction around the
whale's circumference. The two legs
of each harness half (Figure 3) were
attached together ventrally by "D"
rings to a timed-release mechanism.
Dorsally they were bolted to a curved
metal plate holding the radio trans-
mitter. Horizontal rows of grommets
5 cm apart in the heavy plasiic-
impregnated nylon reinforcing band
at the dorsal ends of the harness legs
allowed adjustments to animals of
different circumferences and allowed
the harness to be secured under differ-
ent degrees of tension. We pulled
the harnesses snug on our animals,
which prevented flutter from water
passing around the swimming animal
and kept the harness in place during
dives (the harness was 40 cm wide
and 1 12 cm long).
The strength feature of the harness
was a 2,5 cm wide by 0.6 cm thick
woven nylon strap in the leading and
trailing edge of each harness half.
These straps, held in sewn folds of the
harness, were sewn to the harness only
near the ventral "D" rings, thus per-
mitting harness and straps to be ad-
justed independently to the whale's
circumference. The grommeted ends
of both the harness itself and the
strengthening straps were attached to
bolts on the dorsal plate by means of
knurled nuts.
A plastic cup on each side by mid-
body simulated an instrument housing,
and a poK vinyl chloride rod sewn
across the harness above the pectorals
acted as a batten, preventing bunching
of the harness in the anterior-posterior
direction.
The timed-release mechanism con-
sisted to two aluminum plates held
together by a central spring-loaded
' Reference to Irade names does not imply en-
dorsement by the National Marine Fishieries
Service, NCAA
magnesium bolt. One plate had four
tapered corner posts that tit into four
receptacles on the other plate. The
"D" rings of the harness legs slipped
over the posts and over four strong
springs that assisted in forcing the
plates apart during jettisoning. All
tensions of the harness and nylon
straps were e.xerted against these posts.
The magnesium bolt bore only the
vertical strain of a spring between the
two plates.
The wall thickness of the magnes-
ium bolt determined the interval be-
tween submergence in seawater and
the time of breakage. When the bolt
broke the springs forced the two plates
apart and released the "D" rings from
their posts. Corrosion of the bolts was
insured by a central copper sleeve that
promoted rapid electrolysis.
The dorsal plate to which the har-
ness attached was constructed of ? mm
curved aluminum, designed to fit over
the body contour of a baby whale.
An Ocean Applied Research Model
PT-202 radio transmitter was secured
to the center of this plate, and a paint-
ed yellow cap moulded of high density
polyurethan foam was fitted over the
transmitter for flotation. Foam neo-
prene sheeting was glued to the ventral
surface of this plate to prevent chafing
the whale's skin.
To fasten the harness around the
animal, the two halves, connected
ventrally to the timed-release mechan-
ism, were slid under the animal and
the pectoral flippers inserted through
the harness. The radio, float, and plate
were placed on the dorsal midline,
and each harness half was pulled tight;
the appropriate rows of grommets
in the harness were fitted over bolts in
the dorsal plate and the nuts tightened
down. Then the excess rows of grom-
mets were cut off with a knife and the
heavy nylon straps secured in place
and similarly trimmed. Finally straps
from the float were attached, and the
calf was ready to be launched to its
mother. Figure 4 shows the harness
and radio in place as the released calf
joins its mother.
61
Figure 3. — The harness, radio transmitter and
timed release mechanism: (A) OAR PT-202
radio transmitter (B) 5 mm curved aluminum
dorsal plate (C) Plastic-impregnated nylon re-
inforcing sewn on harness of Lino i^ 241 material
(D) Nylon reinforcing strap sewn into harness
and bolted to dorsal plate (E) Rows of grommets
(F) Knurled nuts holding harness to dorsal plate
(G) Instrument housing (H) Polyvinyl chloride bat-
ten (I) Harness legs with "D" rings (J) Polyurethan
flotation device attached by straps to dorsal
plate (K) Timed-release mechanism (L) Alu-
minum corner posts to which "D" rings of har-
ness legs attach (M) Receptacles for above
posts — spring loaded (N) Magnesium bolt
passes through spring loaded hole in top plate
and secured with a nut.
Dimensions: (1) Harness width 40 cm (2)
Harness length 112 cm from timed — release
mechanism to first row of grommets <3) Distance
between each of five rows of grommets — 5 cm
(4) Length of harness legs 40 cm (5) Timed
release mechanism 10 x is cm.
TRACKING AND
HARNESS RECOVERY
The three harnessed whales re-
mained within a few hundred meters
of their release points (see Figure 1).
Visual tracking in daylight was greatly
assisted by the bright yellow float and
upper harness which were visible even
a foot or two underwater.
in the first release, after some time
in the water, the calf swam purpose-
full> toward the Unison, turned on
her side, and rubbed the harness
against the hull and keel of the boat —
breaking the float partly loose, releas-
ing one "D" ring, and snapping the
fiberglass radio antenna. Transmission
of radio signals immediately ceased.
This damage could have been prevent-
ed by our maintaining a greater dis-
tance from the harnessed animal. The
timed-release mechanism contained a
."^-hour bolt which had not released
by the time darkness fell. The harness
was recovered 2 days later in vegeta-
tion along the channel edge, about 2
km from the release point.
The second release, timed for some-
what less than 5 hours, went flawlessly,
including radio tracking and harness
release.
The third release was planned for
20 hours, with tracking overnight
from the Loiisim. To assist after dark
should the radio malfunction, a water-
proof lifejacket light was ti.xed to the
float. Though both radio and light
functioned at release, thev failed before
62
Figure 4. — Mother and young swimming with harness and radio in place.
dark, and the animal was lost during
the night. However, shortly after dawn
the released harness was found floating
within 60 m of the vessel. Details
of these releases and trackings are
presented in Table 1.
DISCUSSION
The capture methods described here
for suckling gray whales are remark-
ably effective and simple. Except when
the mother is under the pulpit or at
the edge of shallow water, the methods
seem relatively safe. Given enough
shipboard power the noosing methods
would work with larger animals,
though the sheer bulk of an adult
would make any movement by the
whale, purposive or not. dangerous.
This would certainly be a prime con-
sideration in any attempt to affi.x a
harness on an adult.
The harness described here would,
with minor modifications, serve nicely
for short-term tracking and instru-
mentation of small gray whales. Be-
cause a whale attains 66-72 percent of
its adult size in the first year (Rice
and Wolman. 1971). growth during
the first months is e-xtremely rapid.
Harnesses for periods of more than a
week must therefore include a device
that allows for growth but also keeps
a constant tension and locks if the
Table 1. — Harnessing and tracking of gray whale calves.
Animal
Planned
Netting
to
number
Girth
Length
bolt life
beachir
'9
Beaching
to
Date
and name
Sex
(m)
(m)
(hr)
(mm)
release (m
in)
Tracking
1/27/73
(1) Carl(a)
F
2 41
4 80
5±
30 ±
37 ±
4 hr, 45 mm
track, release
time uncertain
1/28/73
(2) Lee
M
2 17
4,20
5±
18 ±
14 ±
3 hr, 16 mm
1/29/73
(3) Baia
F
2.51
5 16
20 ±
28 ±
8±
17 hr. 23 mm
to harness
recovery. Time
to release
uncertain.
animal rubs the harness against under-
water obstructions.
Another concern on any long-term
track is abrasion of the harness. The
purposive rubbing of whale No. 1
against the capture ship and its mother
caused damage to the radio antenna
and serious abrasion to the lower har-
ness legs. On whale No. 2 the abdomi-
nal legs of the harness were abraded
through the girdle fabric and into the
flat nylon strap in several places, even
though the animal wore the harness
for only ?• hours. 16 minutes. Behavi-
oral observations suggest that much
harness wear results when the baby
rubs against the barnacle-covered
back of the mother and slides to one
side as she surfaces. None of these
problems was more than very minor
in these tests. But clearly long-term
tracking with increased exposure to
obstacles along the migratory path
will exacerbate these problems greatly.
More durable materials, such as metal
or the strongest fabric, and more re-
silient radio antennae will be needed
for successful long-term tracking.
The release mechanism dependent
upon magnesium bolt corrosion worked
adequately, but variations in water
temperature and salinity could un-
predictably alter release time. Long
release times (more than a week) may
require a new system, such as the use
of electroexplosive or electronic re-
lease mechanisms that might allow an
operator to release the harness upon
command.
Harnessing is probably the least
injurious means of attaching instru-
ments to cetaceans, and harness place-
ment around the pectoral flipper area
is probably optimal. Pectoral place-
ment insures maximum exposure of
the antenna, minimal body movement
63
during swimming, and relatively little
change in girth during diving. Further,
when physiological data are to be tak-
en, most important vital areas (lungs,
heart, brain) are nearby.
In our opinion package volume
could be relativeh high, providing it is
weight compensated until nearly iso-
static. A baby whale might well carry
15-20 kg of instruments properly
housed and shaped to reduce drag.
Instrument placement is probably best
just above or between pectorals where
it would cause the least disequilibrium.
In these positions it would be most
difficult for the whale to rub the instru-
ments loose. Any such package, of
course, would have to be strongly
protected from impact and abrasion.
The harness used here was designed
with a float at the top to suspend the
antenna with the harness hanging be-
low so that when cast off it rode easily
with the antenna in the vertical posi-
tion for good transmission.
In conclusion, the first steps of whale
capture and instrumentation have been
taken, but much remains to be done
to transfer the methods to (I) long
term trackings. (2) other species which
must be caught and handled at sea,
and {}) adult whales.
Jose Castello of the Consejo Nacional
Ciencias y Tecnologi'a, Mexico City;
and Jaime Dommguez and Mario
Camparan of the Escuela Superior de
Ciencias Marinas, Ensenada, Baja
California Norte, Mexico.
We thank Frank Brocato for con-
sultation on whale capture and for
constructing much of our gear.
We are especially grateful to Edwin
Janss and Richard Wheeler of the
Janss Foundation for support of the
field study. A National Aeronautics
and Space Administration subcontract
No. 23196 (NA.SA NAS2-68601 al-
lowed harness preparation.
LITERATURE CITED
Anonymous. In press. Report of Working
Group on Bio. and Nail. Hist. In W. E.
Schevill (editor). The Whale Problem.
Harvard Press, Cambridge, pp .''-10.
Clarke, R. \^^1. Migrations of marine
mammals. [In Engl, and Norw.] Nor.
Hvalfangsl-Tid. 46:609-6.K).
Cousteau, J. Y. 1972. The whale. Mighty
monarch of the sea. Doubleday &. Co.,
Garden City, .104 p.
Eberhardt. R. L., and K. S. Norns. 1964.
Observations of newborn Pacific gray
whales on Mexican calving grounds. J.
Mammal. 4.S(1):88-9.S.
Rice, D. W.. and A. A. Wolman. 1971.
The life history and ecology of the gray
whale {Exchruiuiiis rohusiti.s). Am. Soc.
Mammal.. Spec. Publ. No. .1, 142 p.
Spencer. M. P. 197.1. Scientific studies on
the gray whales of Laguna Ojo de liebre
(Scammon's Lagoon), Baja California,
Mexico. Natl. Geogr. Soc. Res. Rep., 1966
projects, p. 23-*'-25.1.
MFR Paper 1059. From Marine Fisheries Review, Vol.
36, No. 4, April, 1974. Copies of this paper, in limited
numbers, are available from D83, Technical Information
Division. Environmental Science Information Center.
NOAA, Washington. DC 20235.
ACKNOWLEDGMENTS
Permission to study the gray whale
came from both the Mexican and
United States governments, and many
people helped. Prominent were Carl
L. Hubbs of Scripps Institution of
Oceanography; George Gross, U.S.
Fisheries Attache, U.S. Embassy,
Mexico City; and Philip Roedel and
Robert Miller of National Oceanic
and Atmospheric Administration, U.S.
Department of Commerce. We thank
our willing and skillful field crew:
Captain Tim Houshar and the crew
of the Louson, Richard Pierce, Ken-
neth Balcomb. and Thomas P. Dohl
of the University of California, Santa
Cruz; Gerald Kooyman of Scripps
Institution of Oceanography; Robert
Gibson oi the Franklin Institute Re-
search Laboratories. Philadelphia;
Opposite. — Gigi II, witti transmitter affixed to
back, awaits release at sea off San Diego. Ptioto
by J. S. Leattierwood, courtesy of Naval Under-
sea Center. San Diego, Calif.
Vf GPO 799 418
64
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