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Comparative lipid patterns in acoustical and nonacoustical fatty tissues of dolphins, porpoises and toothed whales
p. 591 to 597. Pergamon Press. Printed in t
PID PATTERNS IN ACC AL FATTY TISSUES OF ES AND TOOTHED WH
Idl
1,11• , I . . ~ I L Y i £ U I I J l I I ~ I l t l
• GREENBERGx, DAVID K. CALDWELL J. C. Sleos s AND R. G. ACKMANs Rutgers University, New Brunswick, N of Florida, St. Augustine, Florida 32084 I ~ l ~ l l l ; l i ~ es Cl, UI*.[ IYIO, I I I I U IdiOt V l l . , ~ l lHalifax, la.lll~k and Marine Service. Nova Sc
L AND INS, CALDWELLs, pmmunication SDepartment B3J 2R3.
(Received 3 April 1974)
analyzed for Abstract--1. Jaw and blubber fats from twenty different ger enera of Od lipid class composition and for iodine value. Results were compared co~ wil ts analyses on melon fats from the same animals. ! iodine value 2. Melon and jaw fats in each animal are very similar in lit)id class co but differ significantly from the corresponding blubber fats. The head : osed of lower molecular weight and more saturated lipids rids than are found found in the th, blubber. Del ~oenidac and 3. High levels of isovalerate lipids are found only in the families fi , longer chain Monodontidae. The Ziphiidae, Physeteridae and Platanistidae Platanistida, fats are length lipids. 4. Delphinidae head fats consist of triglycerides plus signnificant am~ of wax ester. Phocoenidae and Monodontidae head fats are almost exclusively exclusive triglycer or no ( < 3Vo) wax ester present. 5. Ziphiidae blubber fats are almost entirely (>~94~o) wax ester, es a unique characteristic distinlguishing them from all other whales. 6. Relative velocity of sound waves in odontocete head fats fat would be lowest in the families families, Delphinldae, Phocoenldae and Monodontidae, intermediate in the Ziphiidae and highest in the Phy~,,seteridae and Platanistidae. This may indicate several types of echolocation mechanisms withi hin the Odontoceti.
INTRODUCTION
MEMnERS of the suborder Odontoceti (dolphins, porpoises ises. and toothed.. whales). .empl°y. ultrasonic ..... echolocation to navigate and to locate food in their aquatic environment (Norris, 1968, 1969; Evans, 1973). The sound transmitter for this process ges, directly in front appears to lie in the nasal passages, of the skull and behind the large :e fatty melon in the forehead of these animals. Hence orris (1968) has ce Norri Norris postulated that the melon functions ions as an acoustical transducer and sonic lens which1 gives directionality ;holocation. to sound production during echolocati cation Reeeption of the reflected signals apparently involves transmission through the two fat1 fatty bodies located inside the lower mandible (Bullock allock et aL, 1968; Norris, 1968) although experimental aaental evidence for this hypothesis is somewhat incomplete. The unique isovalerate wax esters ters and triglycerides found in several odontocete melon helen and jaw fats (Varanasi & Malins, 1970a, b, 1971, 971, 1972; Litchfield et aL, 1971 a, b; Robisch et al., 197~ ~72) has led Varanasi & Malins (1971, 1972) and Blomber 31omberg (1972) to postulate that isovalerate lipids in particular endow the fatty melon and jaw tissues with special acousti-
docation. However, ,erties favorable for echolocatio cal prope: our li] composiour recent recen comparative survey of the lipid Litchficld & Greentions of codontocete melon fats (Litchfi¢ ber2. 197, berg, 1974) indicated that isovalerate lilpids occur in only three out of the six families in 1the suborder Odontoceti. Apparently more than one type of successfully in melon lipid composition can function st echolocation. We have now extended this inve investigation by making a survey of lipid class composit ~positions (including isovalerate subclasses) and iodine values in the jaw and (for comparison) blubber fats fa of twenty odontocete genera. With these results, result it is now orize the acoustic possible to distinguish and to categorize and non acoustic fatty tissues of the entire en Odontoceti. MATERIALS AND METHOD ETHODS Materials
Most of the fatty tissues used in thi this study were obtained from the same animals as descrit described previously (Litchfield & Greenberg, 1974). Several additional a animals were also sampled: Feresa attenuata (0.82-m juvenile male caught ,,~550 km off the Pacific P~ coast of
591
D. K. CALDWELL,M. C. CALDWELL,J. ,1969); Hyperoal sample from Scotia, Canada, delphis borealis cad near Areata, onoceros jaw fat reals caught in noeeros blubber a Bay, northern 5); Mesoplodon Crescent Beach, aldwell, 1971); (7~Olr(.t~l?l~ ~LII4:ILI~ ~tJ.CJ.LLU~fOL ~LL .L¥]LOAL~O*IIOL JL)~,,O~%,J.I9 Makaha Beach Hawaii, November 1967). Where possible, jaw s were extracted from the total mandibular fat ad the blubber was sampled near the dorsal fin.
Delphinidae jaw predominant subcl Pseudorca blubber this sample was ta adjacent to the mel
KV species as the XX wax ester in Lbly not typical, for abber tissue directly :riglycerides, VXV is l the jaw fats, while tjor (Lagenorhynchus blubber fats. This be concentrated in clearly illustrated by lain moieties in the ale 2). Delphinidae, e melon and jaw fats ~ile the blubber fats O0%). s (Ziphiidae, Physeag-chain lipids were elations exist within
the predominant sl XXX is the predom and Phocoena) su~ tendency for isova the melon and jaw calculating the mc [o[a[ llpF. of each total lipids Phocoeni aidae and 1~ contain 338-59~ 1ol possess aossess n~ much highe In the nonisoval Is teridae, Platanistid l Is were extracted, hydrogenated and separated by found; and a] compo ~t not fan Rive thin-layer chromatography (TLC) as des- but not between bq ares of triglycerides Ziphiidae jaw f~ in our previous paper (Litchfield & Greenberg, Ziphiid small amounts of Each sample contained one to five major (I> 1-2~) and wax esters t( (triglycerides, diacyl glyeeryl ethers, wax esters) diacyl glyceryl gl are similar to the etl a,ere recovered and quantitated by measuring the Berardius melon fat Lpreviously reported ~rption of the ester linkage at 1745 cm -a (Freeman, (Litchfieh Litchfield & Greei On the other hand, Atchfield & Greenberg, 1974). As noted before the Zipl~ )hiidae blu tre almost entirely eld et al., 1971a; Litchfield & Greenberg, 1974), (~>94~)) long-chain 1 I unique characterissenee of isovalerie acid in such molecules permits distinguishing the Ziphiidae fro] from all other C separation of lipid subclasses according to the tic distir whales. 11 LII l l i . / ~ ; l: of long-chain(X) and isovaleroyl (V) moieties in the moleeul Leeule. The wax esters are resolved into isovaleroyl In the Physeteridae, there are only two genera, ( x v ) and aad nonisovaleroyl (XX) bands; and the trigly- both of which have been examined. All melon cerides separate into three bands (XXX, XXV, VXV) (Litchfieh Litchfield & Gre~nberg, 1974; Hanse Hansen & Cheah, containin dng 0, 1 and 2 isovalerate moieties. Any isobuty- 1969), jaaw and blubber fat sample )les contain a rate or 2-methylbutyrate esters, which are occasionally mixture of c long-chain wax esters and triglycerides; present in minor amounts in odontocete fats (Nakamura but the ccompositions of fats from acoustical ac and & Tsujino, ino, 1952; Schlenk, 1965; Morii & Kanazu, 1972; nonacoustical nonacous tissues are not clearly differentiated. d Ackmann e t al., 1973), would act the same as isovalerate In v i e w of recent findings that lipid composition esters durin uring TLC and would be reported as "V". widely within the large melons o f Physeter The iodine value of each tissue fat was determined varies wi . . . . . . . . . . . . . . . . . . . . . . {l~/[r~rP;e ,itchfield & Karol, mericanOilCbemists' (Morris, 1973) and Kogia (Litchfieh using the standard Wijs method (American ( ( an Oil unpublished), the question arises whether the ' Society, 1967) scaled down to 10 mg~,size s~ze samples, samples analyzed here represent typical typi~ compositions for their respective tissues. Onlyt further study RESULTS AND DISCUSSION can clarify this matter. In the Platanistidae, lipid class compositions Compositional analyses apparently vary widely from genus to genus. Inia The lipid class compositions of odontocete jaw contains a mixture of long-chain wax wa esters and and blubber fats reported in Table )le 1 show a number triglycerides in the melon (Litchfield & Greenberg, of clearcut correlations with taxonom bnomy and anatomy. 1974) and jaw fats, but only triglyc~ ,cerides in the Animals in the families Delphinidae, [nidae, Phocoenidae blubber. Platanista gangetica, however lowever, possesses a and Monodontidae all contain subst~ substantial amounts melon fat consisting almost exclusive xclusively of triglyof isovalerate (V) lipids, while the Ziphiidae, cerides (Litchfield & Greenberg, 19] 1974), and the Physeteridae and Platanistidae do not. There are blubber fat is apparently similar (Pathal Pathak et al., 1956; distinct compositional differences ~s between the jaw Tsuyuki & Itoh, 1971). and blubber lipids in all families ies, but the jaw fat The relative unsaturation of odontq odontocete melon, compositions closely resemble the corresponding jaw and blubber fats was also studied b~ determining melon fats previously analyzed (iLitchfield & Green- the iodine values of all available saml ~les. Results berg, 1974). (Table 3) indicate that the melon and jaw fats are Within the isovalerate-containin ining families (Del- markedly more saturated than the corresponding c~ phinidae, Phocoenidae, Monodontid,' Lodontidae), trigly- blubber fats, except for Kogia (see remark rem~ above on cerides are the major lipid class present. ~resent. Appreciable Kogia samples). In the Delphinidae, bl blubber iodine amounts of wax esters (3--49~)) occur only in the values are 78-125 units higher than the t respective
rid patterns in odontocete fatty tissues osition of odontocete jaw and blubber fi Jaw aeyl
~eryl lers
Triglycerides XXX XXV VXV
1pus griseus 1 norhynchus albirostris delphis borealis 1 torca crassidens 3 riafluviatilist 1 ;llaplagiodon 4 'ops truncatus 12 :nidae oena phocoena oenoides dalli
5 2 16 48 41 28
--
----
<1 <1
[ontidae hinapterus leucas~ 9don monoceros ae rdius bairdi 36 - Hyperoodon ampullatus Mesoplodon densirostris Ziphius cavirostris 19 - Physeteridae Kogia simus Physeter catodon§ Platanistidae Iniageoffrensis
18 - -
66 - -
1
7
20
67
14 2 Tr
28 7 3
Triglycerides
Wax ester., XX
XXX XXV VXV
XV
11 15 7 43
---16
96
--
53
40
1
3
55 72 48 51
3
5
52
83 68
17 32
-7 ---
10 20
35 38
54 42
29 58
41 36
30 6
19
33
48
74 95
21 2
5 --
63
--
--
- -
- -
--
--
--
--
Tr
4 ---
97
--
94 - 99 - 2
79
--
--
1
81
--
--
2
89 85 93 41
32
--
--
42
--
79
--
6
Tr
58 21 99
* X, Long-chain (~>Ca) fatty alcohol or fatty acid. V, Isovaleric acid. Order of letters in triglyt ceride abbreviation does nott indicate :ate an any specific isomer composition. t From Litchfieid etal. (1971b). $ From Litchtield et al. (1971a). § From Hansen & Cheah (1969). ~tocete families, this head fats! In the other five odontocete te families fi difference is 23-89 units (except for or Kogia Kogia). )eia). We have compared the anallyses reported here with published data on lipid classs composition, fatty acid composition a n d iodine value value of fats from Delphinus (Schlenk, 1965), Stenella ~tenella (Morii & Kanazu, 1972), Tursiops (Gill & Tucker, 1930; Varanasi & Malins, 1970a, 1971, q , 1972), Phocoena (Lovern, 1934; Varanasi & Malins. ins, 1970b), Phocoenoides ( N a k a m u r a & Tsujino, 1952; T a k a o k a & Tsujino, 1952), Monodon (Robisch )isch et aL, 1972), soplodon (Andre & Berardius (Saiki, 1953a, b), Mesoplodon Canal, 1926), Ziphius (Igarashi ett aL, al. 1953), Physeter (Morris, 1973), Inia ( A c k m a n et al., 1971) a n d Platanista (Pathak et aL, 1956 ; Tsuyuki & Itoh, 1971, 1972). Overall agreement is fairly good, even though the exact compositions) reported are not
always identical with our findings for the same genus, especially with respect to the a m o u n t of isovaleric acid present. Such differences probabl'. )robably arise from (i) the usual inter- a n d intraspecies variations vari| in the lipid composition of animals, (ii) the varying anatomical locations from which fatty tissues were sampled, (iii) the wide variations in isovalerate animal content found within the melon of a single s (Litchfield et al., 1973; W e d m i d et al., al. 1973), (iv) our selection of only t> 1-2% T L C hands b a n (i.e. those of acoustical importance) for quantita uantitation and ( v ) differences in analytical techniques. I n almost all eases, however, there is i agreement between the literature data a n d the comparative lipid patterns which we have found. found Apparent exceptions are the high iodine values reported by lla mel¢ melon (44.5 I.V.) Morii & K a n a z u (1972) for Stenella
D. K. CALDWELL,M. C. CALDWELL,J. in odontocete its
Monodontidae util major portion of th~
Nay for producing a 5.
Jaw:~ Blubber:~ Implications for ech, 96 94 43
97 71
--'~".... torca !ia ;lla 'ops
44 38 41 43
49 40 43 43 47
100 79 93 88
nidae oena oenoides
41 59
48 55
62 81
[ontidae hinapterus odon
38 46
52
87 99
ae rdius ,roodon ~plodon ~us Physeteridae Kogia Physeter Platanistidae Inia Platanista
100
100
100 100 1O0
100 1O0 1O0 100 100
100
100
100 100 100 100
ng-chain acids and alcohols: X>~Ca. Short-chain * Lon V
Analytical result, berg, 1974) show th melon and jaw fat ponding blubber f~ three genera for implies that the ev fat compositions h~ Iat comp~ specific functional f fats have quite sin: [y that likely tha they are (1968 tion. Norris No and fatty melon me transduce for the transducers
; Litchfield & Greensitions of odontocete tly from the corres-two of the twentyare available. This lese distinctive head :tive advantage for a the melon and jaw aapositions, it seems ,~d in the same funcLted that odontocete serve as acoustical used in echolocation.
valt Table 3. Iodine I
cete melon, jaw and
Family Genus
on
Jaw
Delphinidae Delphinus Feresa Globicephala Grampus Lagenorhynchus Lissodelphis Pseudorca Sotalia Stenella Tursiops
10 6 20 8 17 6 5 6
Phocoenidae Neophocaena Phocoena Phocoenoides
44* 24 53
39 41
Monodontidae Delphinapterus Monodon
23 30
34
18
7
Ziphiidae Berardius Hyperoodon Mesoplodon Ziphius Physeteridae Kogia Physeter
Blubber 140 92
21 35 9 11
6 17
131 99 106 89 121 109 87* 86 79 112 90 82 86 76
14 69 541-
76
76 91t
Platanistidae 25 73 Inia 31 99~ Platanista 76 * From Kanazu et aL (1969). Neopho ~hocaena phocaeScheffer, 1968). noides = Neomeris phocaenoides (Rice & S l" Calculated from Hansen & Cheah (196! C1969). ~:From Pathak et al. (1956).
pid patterns in odontocete fatty tissues heir distinctive > 50 mole %) of id as isovaleric on and jaw fats }72) and Biotarate wax esters tical properties ch are essential point out that mtly lower in ~rate npms man m me corresponamg ;ponding ranglong(i.e. blubber fat) compounds (Gouw & :r, 1967). Our comparative compositional data te, however, that no single lipid class, lipid ss or isovalerate composition is uniquely tted with echolocation ability in the Odontothis lack of echolocation]isovalerate correlaoes not necessarily invalidate the Varanasis-Blomberg hypothesis, however, It means the Delphinidae/Phocoenidae/Monodontidae merely represent a special case, for there are ely other melon and jaw lipid compositions can successfully function in echolocation, then raises the question of how many dis'e types of lipid composition can be identified mtocete melon and jaw fats. At the present ve can distinguish four groups (Fig. 1). Delphinidae ae head fats contain isovaleric acid, triglycerides and significant amounts (>I 3%) of wax ester, Odonfocefi
Isovoleric
No isovaleric
~d > 3 0 wax
esfers Delphinidae
wax esters
mostly C~o-ci2
Phocoenidae Z~ol~idae Monodontidae
¢q0"C20 oeids
I
Physe~ridae Plafanisfidae
Fig. 1. Comparative lipid patterns in odontocete melon and jaw fats based on analyses of nineteen of the thirtythree living genera. Phocoenidae and Monodontid~[ae head fats also contain isovaleric acid and trigbycerides but little or no (~< 3%) wax ester. Ziphiidae head fa fats possess no isovaleric acid and no distinctive li lipid class composttion; but work now in progresss (Litchfield et al., unpublished) and early distillation lation analyses by Saiki (1953a) indicate that all Zi~iphiidae melon and jaw fats contain large amounts of['C,o--C~fattyacids C not found elsewhere in the Odon toceti. Physeteridae and Platanistidae head lipids possess )ssess no significant amount of isovaleric acid nor av.ly known distinctive lipid class or fatW acid composition.
Since it is the ac erties of odontocete head fats that wou ant in echolocation, compositional data ably be classified on this basis. The v, )und in lipids is a regular function of lax weight (Gouw & Vlugter, 1967) and with the isovaleric acid, C,0-C,s acic chain acid content. On this basis, odol fats can be divided into low (Delphinid ]ae, Monodontidae), intermediate (Zip11 high (Physeteridae, Platanistidae) tool. it categories, each possessmj charact| possessing velocities. Rcpresentative triglyceri (trioctanoin, Cz4), intermediate (trilau intermedi high (triolein, Cs4) carbon numbers n c ~Iz sound waves at 1315, 13: 1357 and 1 ~spectively at 40°C (Gouw & Vlugter, :perimental data are available on sound wax esters, but the same trend tre~ of inct ~y at higher carbon numbers,would be e numbers Can these th three ]ontoceti possessing different acoustical a their head fats be correlatexd with thr ; types of echolocation mechanisms mecln ?( ace is too incomplete to answe wet this qu there is speculative evidence pointing 1 iF n. Norris & Harvey (1972) have ha recently proposed a new t: type of ultrasonic transmission tra for echolocation purposes in Physeter catodon; this involves reflection reflec off the posterior surface of the spermaceti cas~e rather than the unidirectional unidi forward transmissh )n found in dolphins (Norris, 1968; Evans, 1973). The unique convoluti convolutions of the rostrum and mel on in Kogia (Schenkk Schenkkan & Purves, 1973) also suggG:est a distinctive u r nasonic transmission mechani :hanism in that Moreover, Ziphiidae melon tissue tiss structures animal. 1~ and rostr ostrum shapes are known to differ greatly from the corresponding anatomy of the Delphinidae, 1 Phocoenidae, and Monodontidae (N (Norris, 1964; Litchfield & Greenberg, unpublished) blisbed). However, only further study can reveal whether the t~ distinctive head fat compositions of the Ziphiidae, hiidae, Physeteridae and Platanistidae are related to theirr unusual unu cephalic anatomies and their mechanisms of echolocation. echo
Acknowledgements~We extend our th thanks to the following for their help in securing o¢ odontocete fat samples for us in this study: Marineland • ineland of Florida; Dr. E. D. Mitchell, Dr. A. W. Mansfield ant Mr. B. Beck ~field and (Fisheries and Marine Service, Ste. Anne de Bellevue, Canada); Dr. H. Omura and Dr. A. Kawar Kawamura (Whales Research Institute, Tokyo); Dr. P. B. Best (Director of Sea Fisheries, Cape Town); Dr. R. P. van va der Merwe (Fishing Industry Research Institute, Rondebosch, South Africa); Dr. J. G. Mead (Smithsonia fithsonian Institution, University College of Washington, D.C.); Dr. I. Rees (Universi North Wales, Menai Bridge); Dr. H. Olcott Oleo (University of California, Davis); Dr. D. C. Malins and Dr. A. Wohiman (National Marine Fisheries Sela Service, Seattle); Dr. K. S. Norris (University of California, Santa Cruz); Dr. W. J. Houck (California State Univer,sity, Arcata);
D. K. CALDWELL, M. C. CALDWELL, J. eries Service, La x, Canada); and hill, Manitoba). ynthesis of the yses. part by grants (No. GB-30553) ncil.
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LITCHFIELDC. & Gr I. (19741 Comparative lipid patterns inthe olphins, porpoisesand toothed whales. Cc Physiol. 47B, 401-407, LITCHFIELD C., KAR EENBERG A. J. (1973) Compositional tol melon lipids in the Atlantic bottleno Tursiops truncatus: implications for ecl dar. Biol. 23, 165-169. LITCHFIELD C., KIN/~ :MAN R. G. & EATON C. A. (1971b) C pid patterns in two freshwater dolphi ffrensis and Sotalia fluviatilis. J. Am. C 48, 91A. abolism in fishes--Ill. LOVERNJ. A. (1934) ots. Biochem. J. 28, Selective ,.3t:lectlvt formatio 394-402. KANAZE tty acids in the foetus, MORIIH. nurselin g and adt of dolphin, Stenella attenuata. 7ish. 38, 599-605. attenuat Bull. Jal MORRIS ture of the spermaceti MORRISR. R J. (1973)" seter catodon). Deeporgan of o the spen Sea Res. Res 20, 911-91 NAKAMURA 2) The acidic constiNAKAMUR Y. & T: ¢ dalli True) head oil. tuents cof porpoise ,ric. Chem. Sot ),-645. J. Agric ms of echolocation in NORRIS K. K S. (1964) cetaceans. In coustics (Edited by TAVOLGA -336. Pergamon Press, TAVOLG W. N.), "~ New York. f acoustic mechanisms NORRIS K. S. (1968) ~I ,dontocete ceta( rution and Environment in odon Y University (Edited by DRAKE E. T.), pp. 297-324. Yale Press, New Haven. NORRIS K. K S. (1969) The echolocation of marine maremals. In II The Biology of Marine Mamm, Mammals (Edited by ANDERSEN H. T.), pp. 391-423. Academic Acaderr Press, New ANDERS York. NORRIS K. K S. & HARVEY G. W. (1972) A theory for the function functior of the spermaceti organ of the sperm whale (Physeter catodon L.). In Animal Orientation and Navigation (Edited by GALLER S. R., SCHMIDTI; E,, pp. 397KOENIGK., JACOBSG. J. & BELLEVILLE R. 417. N.A.S.A., Washington, D.C. GARWAL C. V. (1956) PATHAK S. P., SUWAL P. N. & AGARWAI. liver fats. Component acids of suinsh blubber and a Biochem. J. 62, 634--637. on a PERRINW. F. & HtmBS C. L. (1969) Observations Obs( young pygmy killer whale (Eeresa attenuata attenua Gray) from the eastern tropical Pacific Ocean. Trans. Tra San Diego Sue. Nat. Hist. 15, 297-308. RICE D. W. & SCHEFFER V. B. (1968) A 1List of Marine Mammals of the World. Special Scientific Scien Report-Service, Fisheries No. 579, U.S. Fish and Wildlife Wi Washington, D.C. ROBISCH P. A., MALINS D. C., BEST R. (~ & VARANASI U. acoustic (1972) Differences in triacylglycerols from 1 tissues and posterior cranial blubber of o the narwhal (Monodon monoceros). Biochem.J. 130, 33P-34P. 3 SAIKI M. (1953a) Studies on whale,- oil--VII. oil--V Fatty acid composition of bottlenose whale oil--I. On the head and jaw oil. Bull. Jap. Soc. scient. Fish. 18, 1! 497-501. oil--VIII. Fatty acid SAIKIM. (1953b) Studies on whale oiI--VI On the composition of Pacific beaked whale oil--2. o blubber oil. Bull. Jap. Soc. scient. Fish. 19, 1! 809-812. 19731 The SCHENKKANE. J. & PURVESP. E. (1973) Th comparative of the anatomy of the nasal tract and the function f~ ~eteridae (Manmaalia, spermaceti organ in the Physeteridae ferkunde 43, 93-112. Odontoceti). Bijdragen Tot de Dierkunde
~id patterns in odontocete fatty tissues analysis of optiChem. Soc. 42, on the body oils rrue landed in 3f body oils and Soc. scient. Fish.
t components of 7Vhales Res. Inst.
)u~
vtuuu~wz
vii
vt
~Jo.115~,o
d component of Sci. *Jut.
x-~tvv,,x r dolphin. t.tVlVllltt.
Whales Res. Inst. 24, 11%125. gI H. & ITOH S. (1973) Fatty acid components of her oil of the Amazon River dolphin. ScL Rep. les Res. Inst. 25, 293-299. ASI U. & MALINS D. C. (1970a) Unusual wax s from the mandibular canal of the porpoise flops gilli). Biochemistry9, 3629-3631.
VARANASXU. & MAt linked lipids in th (Phocoena phocoen( glycerolipids. Bioc~ VARANASI U. (~ MALl
porpoise (Tursiops, and wax esters o melon) and blubb( 231, 415-418. VARANASIU. & MA characteristic of p structures of diiso 176, 926-928. WEDMm Y., Y LrrcHF~ MITCHELL MITcHEJ E. D. (1 the sition within ss whale. Biochim. bi(
Word Index Key W porpoises; whales.
70b) Ester and ethercanal of a porpoise e of isovaleric acid in 6-4579. [) Unique lipids of the :es in triacyl glycerols ~ndibular canal and ochem, biophys. Acta L972) Triacylglycerols tic tissues: molecular ides. Science, Wash. N R. G., Sreos J. C. & ;neity of lipid comport tissue of the pilot 5, 439-447. ty tissue; dolphins;
PID PATTERNS IN ACC AL FATTY TISSUES OF ES AND TOOTHED WH
Idl
1,11• , I . . ~ I L Y i £ U I I J l I I ~ I l t l
• GREENBERGx, DAVID K. CALDWELL J. C. Sleos s AND R. G. ACKMANs Rutgers University, New Brunswick, N of Florida, St. Augustine, Florida 32084 I ~ l ~ l l l ; l i ~ es Cl, UI*.[ IYIO, I I I I U IdiOt V l l . , ~ l lHalifax, la.lll~k and Marine Service. Nova Sc
L AND INS, CALDWELLs, pmmunication SDepartment B3J 2R3.
(Received 3 April 1974)
analyzed for Abstract--1. Jaw and blubber fats from twenty different ger enera of Od lipid class composition and for iodine value. Results were compared co~ wil ts analyses on melon fats from the same animals. ! iodine value 2. Melon and jaw fats in each animal are very similar in lit)id class co but differ significantly from the corresponding blubber fats. The head : osed of lower molecular weight and more saturated lipids rids than are found found in the th, blubber. Del ~oenidac and 3. High levels of isovalerate lipids are found only in the families fi , longer chain Monodontidae. The Ziphiidae, Physeteridae and Platanistidae Platanistida, fats are length lipids. 4. Delphinidae head fats consist of triglycerides plus signnificant am~ of wax ester. Phocoenidae and Monodontidae head fats are almost exclusively exclusive triglycer or no ( < 3Vo) wax ester present. 5. Ziphiidae blubber fats are almost entirely (>~94~o) wax ester, es a unique characteristic distinlguishing them from all other whales. 6. Relative velocity of sound waves in odontocete head fats fat would be lowest in the families families, Delphinldae, Phocoenldae and Monodontidae, intermediate in the Ziphiidae and highest in the Phy~,,seteridae and Platanistidae. This may indicate several types of echolocation mechanisms withi hin the Odontoceti.
INTRODUCTION
MEMnERS of the suborder Odontoceti (dolphins, porpoises ises. and toothed.. whales). .empl°y. ultrasonic ..... echolocation to navigate and to locate food in their aquatic environment (Norris, 1968, 1969; Evans, 1973). The sound transmitter for this process ges, directly in front appears to lie in the nasal passages, of the skull and behind the large :e fatty melon in the forehead of these animals. Hence orris (1968) has ce Norri Norris postulated that the melon functions ions as an acoustical transducer and sonic lens which1 gives directionality ;holocation. to sound production during echolocati cation Reeeption of the reflected signals apparently involves transmission through the two fat1 fatty bodies located inside the lower mandible (Bullock allock et aL, 1968; Norris, 1968) although experimental aaental evidence for this hypothesis is somewhat incomplete. The unique isovalerate wax esters ters and triglycerides found in several odontocete melon helen and jaw fats (Varanasi & Malins, 1970a, b, 1971, 971, 1972; Litchfield et aL, 1971 a, b; Robisch et al., 197~ ~72) has led Varanasi & Malins (1971, 1972) and Blomber 31omberg (1972) to postulate that isovalerate lipids in particular endow the fatty melon and jaw tissues with special acousti-
docation. However, ,erties favorable for echolocatio cal prope: our li] composiour recent recen comparative survey of the lipid Litchficld & Greentions of codontocete melon fats (Litchfi¢ ber2. 197, berg, 1974) indicated that isovalerate lilpids occur in only three out of the six families in 1the suborder Odontoceti. Apparently more than one type of successfully in melon lipid composition can function st echolocation. We have now extended this inve investigation by making a survey of lipid class composit ~positions (including isovalerate subclasses) and iodine values in the jaw and (for comparison) blubber fats fa of twenty odontocete genera. With these results, result it is now orize the acoustic possible to distinguish and to categorize and non acoustic fatty tissues of the entire en Odontoceti. MATERIALS AND METHOD ETHODS Materials
Most of the fatty tissues used in thi this study were obtained from the same animals as descrit described previously (Litchfield & Greenberg, 1974). Several additional a animals were also sampled: Feresa attenuata (0.82-m juvenile male caught ,,~550 km off the Pacific P~ coast of
591
D. K. CALDWELL,M. C. CALDWELL,J. ,1969); Hyperoal sample from Scotia, Canada, delphis borealis cad near Areata, onoceros jaw fat reals caught in noeeros blubber a Bay, northern 5); Mesoplodon Crescent Beach, aldwell, 1971); (7~Olr(.t~l?l~ ~LII4:ILI~ ~tJ.CJ.LLU~fOL ~LL .L¥]LOAL~O*IIOL JL)~,,O~%,J.I9 Makaha Beach Hawaii, November 1967). Where possible, jaw s were extracted from the total mandibular fat ad the blubber was sampled near the dorsal fin.
Delphinidae jaw predominant subcl Pseudorca blubber this sample was ta adjacent to the mel
KV species as the XX wax ester in Lbly not typical, for abber tissue directly :riglycerides, VXV is l the jaw fats, while tjor (Lagenorhynchus blubber fats. This be concentrated in clearly illustrated by lain moieties in the ale 2). Delphinidae, e melon and jaw fats ~ile the blubber fats O0%). s (Ziphiidae, Physeag-chain lipids were elations exist within
the predominant sl XXX is the predom and Phocoena) su~ tendency for isova the melon and jaw calculating the mc [o[a[ llpF. of each total lipids Phocoeni aidae and 1~ contain 338-59~ 1ol possess aossess n~ much highe In the nonisoval Is teridae, Platanistid l Is were extracted, hydrogenated and separated by found; and a] compo ~t not fan Rive thin-layer chromatography (TLC) as des- but not between bq ares of triglycerides Ziphiidae jaw f~ in our previous paper (Litchfield & Greenberg, Ziphiid small amounts of Each sample contained one to five major (I> 1-2~) and wax esters t( (triglycerides, diacyl glyeeryl ethers, wax esters) diacyl glyceryl gl are similar to the etl a,ere recovered and quantitated by measuring the Berardius melon fat Lpreviously reported ~rption of the ester linkage at 1745 cm -a (Freeman, (Litchfieh Litchfield & Greei On the other hand, Atchfield & Greenberg, 1974). As noted before the Zipl~ )hiidae blu tre almost entirely eld et al., 1971a; Litchfield & Greenberg, 1974), (~>94~)) long-chain 1 I unique characterissenee of isovalerie acid in such molecules permits distinguishing the Ziphiidae fro] from all other C separation of lipid subclasses according to the tic distir whales. 11 LII l l i . / ~ ; l: of long-chain(X) and isovaleroyl (V) moieties in the moleeul Leeule. The wax esters are resolved into isovaleroyl In the Physeteridae, there are only two genera, ( x v ) and aad nonisovaleroyl (XX) bands; and the trigly- both of which have been examined. All melon cerides separate into three bands (XXX, XXV, VXV) (Litchfieh Litchfield & Gre~nberg, 1974; Hanse Hansen & Cheah, containin dng 0, 1 and 2 isovalerate moieties. Any isobuty- 1969), jaaw and blubber fat sample )les contain a rate or 2-methylbutyrate esters, which are occasionally mixture of c long-chain wax esters and triglycerides; present in minor amounts in odontocete fats (Nakamura but the ccompositions of fats from acoustical ac and & Tsujino, ino, 1952; Schlenk, 1965; Morii & Kanazu, 1972; nonacoustical nonacous tissues are not clearly differentiated. d Ackmann e t al., 1973), would act the same as isovalerate In v i e w of recent findings that lipid composition esters durin uring TLC and would be reported as "V". widely within the large melons o f Physeter The iodine value of each tissue fat was determined varies wi . . . . . . . . . . . . . . . . . . . . . . {l~/[r~rP;e ,itchfield & Karol, mericanOilCbemists' (Morris, 1973) and Kogia (Litchfieh using the standard Wijs method (American ( ( an Oil unpublished), the question arises whether the ' Society, 1967) scaled down to 10 mg~,size s~ze samples, samples analyzed here represent typical typi~ compositions for their respective tissues. Onlyt further study RESULTS AND DISCUSSION can clarify this matter. In the Platanistidae, lipid class compositions Compositional analyses apparently vary widely from genus to genus. Inia The lipid class compositions of odontocete jaw contains a mixture of long-chain wax wa esters and and blubber fats reported in Table )le 1 show a number triglycerides in the melon (Litchfield & Greenberg, of clearcut correlations with taxonom bnomy and anatomy. 1974) and jaw fats, but only triglyc~ ,cerides in the Animals in the families Delphinidae, [nidae, Phocoenidae blubber. Platanista gangetica, however lowever, possesses a and Monodontidae all contain subst~ substantial amounts melon fat consisting almost exclusive xclusively of triglyof isovalerate (V) lipids, while the Ziphiidae, cerides (Litchfield & Greenberg, 19] 1974), and the Physeteridae and Platanistidae do not. There are blubber fat is apparently similar (Pathal Pathak et al., 1956; distinct compositional differences ~s between the jaw Tsuyuki & Itoh, 1971). and blubber lipids in all families ies, but the jaw fat The relative unsaturation of odontq odontocete melon, compositions closely resemble the corresponding jaw and blubber fats was also studied b~ determining melon fats previously analyzed (iLitchfield & Green- the iodine values of all available saml ~les. Results berg, 1974). (Table 3) indicate that the melon and jaw fats are Within the isovalerate-containin ining families (Del- markedly more saturated than the corresponding c~ phinidae, Phocoenidae, Monodontid,' Lodontidae), trigly- blubber fats, except for Kogia (see remark rem~ above on cerides are the major lipid class present. ~resent. Appreciable Kogia samples). In the Delphinidae, bl blubber iodine amounts of wax esters (3--49~)) occur only in the values are 78-125 units higher than the t respective
rid patterns in odontocete fatty tissues osition of odontocete jaw and blubber fi Jaw aeyl
~eryl lers
Triglycerides XXX XXV VXV
1pus griseus 1 norhynchus albirostris delphis borealis 1 torca crassidens 3 riafluviatilist 1 ;llaplagiodon 4 'ops truncatus 12 :nidae oena phocoena oenoides dalli
5 2 16 48 41 28
--
----
<1 <1
[ontidae hinapterus leucas~ 9don monoceros ae rdius bairdi 36 - Hyperoodon ampullatus Mesoplodon densirostris Ziphius cavirostris 19 - Physeteridae Kogia simus Physeter catodon§ Platanistidae Iniageoffrensis
18 - -
66 - -
1
7
20
67
14 2 Tr
28 7 3
Triglycerides
Wax ester., XX
XXX XXV VXV
XV
11 15 7 43
---16
96
--
53
40
1
3
55 72 48 51
3
5
52
83 68
17 32
-7 ---
10 20
35 38
54 42
29 58
41 36
30 6
19
33
48
74 95
21 2
5 --
63
--
--
- -
- -
--
--
--
--
Tr
4 ---
97
--
94 - 99 - 2
79
--
--
1
81
--
--
2
89 85 93 41
32
--
--
42
--
79
--
6
Tr
58 21 99
* X, Long-chain (~>Ca) fatty alcohol or fatty acid. V, Isovaleric acid. Order of letters in triglyt ceride abbreviation does nott indicate :ate an any specific isomer composition. t From Litchfieid etal. (1971b). $ From Litchtield et al. (1971a). § From Hansen & Cheah (1969). ~tocete families, this head fats! In the other five odontocete te families fi difference is 23-89 units (except for or Kogia Kogia). )eia). We have compared the anallyses reported here with published data on lipid classs composition, fatty acid composition a n d iodine value value of fats from Delphinus (Schlenk, 1965), Stenella ~tenella (Morii & Kanazu, 1972), Tursiops (Gill & Tucker, 1930; Varanasi & Malins, 1970a, 1971, q , 1972), Phocoena (Lovern, 1934; Varanasi & Malins. ins, 1970b), Phocoenoides ( N a k a m u r a & Tsujino, 1952; T a k a o k a & Tsujino, 1952), Monodon (Robisch )isch et aL, 1972), soplodon (Andre & Berardius (Saiki, 1953a, b), Mesoplodon Canal, 1926), Ziphius (Igarashi ett aL, al. 1953), Physeter (Morris, 1973), Inia ( A c k m a n et al., 1971) a n d Platanista (Pathak et aL, 1956 ; Tsuyuki & Itoh, 1971, 1972). Overall agreement is fairly good, even though the exact compositions) reported are not
always identical with our findings for the same genus, especially with respect to the a m o u n t of isovaleric acid present. Such differences probabl'. )robably arise from (i) the usual inter- a n d intraspecies variations vari| in the lipid composition of animals, (ii) the varying anatomical locations from which fatty tissues were sampled, (iii) the wide variations in isovalerate animal content found within the melon of a single s (Litchfield et al., 1973; W e d m i d et al., al. 1973), (iv) our selection of only t> 1-2% T L C hands b a n (i.e. those of acoustical importance) for quantita uantitation and ( v ) differences in analytical techniques. I n almost all eases, however, there is i agreement between the literature data a n d the comparative lipid patterns which we have found. found Apparent exceptions are the high iodine values reported by lla mel¢ melon (44.5 I.V.) Morii & K a n a z u (1972) for Stenella
D. K. CALDWELL,M. C. CALDWELL,J. in odontocete its
Monodontidae util major portion of th~
Nay for producing a 5.
Jaw:~ Blubber:~ Implications for ech, 96 94 43
97 71
--'~".... torca !ia ;lla 'ops
44 38 41 43
49 40 43 43 47
100 79 93 88
nidae oena oenoides
41 59
48 55
62 81
[ontidae hinapterus odon
38 46
52
87 99
ae rdius ,roodon ~plodon ~us Physeteridae Kogia Physeter Platanistidae Inia Platanista
100
100
100 100 1O0
100 1O0 1O0 100 100
100
100
100 100 100 100
ng-chain acids and alcohols: X>~Ca. Short-chain * Lon V
Analytical result, berg, 1974) show th melon and jaw fat ponding blubber f~ three genera for implies that the ev fat compositions h~ Iat comp~ specific functional f fats have quite sin: [y that likely tha they are (1968 tion. Norris No and fatty melon me transduce for the transducers
; Litchfield & Greensitions of odontocete tly from the corres-two of the twentyare available. This lese distinctive head :tive advantage for a the melon and jaw aapositions, it seems ,~d in the same funcLted that odontocete serve as acoustical used in echolocation.
valt Table 3. Iodine I
cete melon, jaw and
Family Genus
on
Jaw
Delphinidae Delphinus Feresa Globicephala Grampus Lagenorhynchus Lissodelphis Pseudorca Sotalia Stenella Tursiops
10 6 20 8 17 6 5 6
Phocoenidae Neophocaena Phocoena Phocoenoides
44* 24 53
39 41
Monodontidae Delphinapterus Monodon
23 30
34
18
7
Ziphiidae Berardius Hyperoodon Mesoplodon Ziphius Physeteridae Kogia Physeter
Blubber 140 92
21 35 9 11
6 17
131 99 106 89 121 109 87* 86 79 112 90 82 86 76
14 69 541-
76
76 91t
Platanistidae 25 73 Inia 31 99~ Platanista 76 * From Kanazu et aL (1969). Neopho ~hocaena phocaeScheffer, 1968). noides = Neomeris phocaenoides (Rice & S l" Calculated from Hansen & Cheah (196! C1969). ~:From Pathak et al. (1956).
pid patterns in odontocete fatty tissues heir distinctive > 50 mole %) of id as isovaleric on and jaw fats }72) and Biotarate wax esters tical properties ch are essential point out that mtly lower in ~rate npms man m me corresponamg ;ponding ranglong(i.e. blubber fat) compounds (Gouw & :r, 1967). Our comparative compositional data te, however, that no single lipid class, lipid ss or isovalerate composition is uniquely tted with echolocation ability in the Odontothis lack of echolocation]isovalerate correlaoes not necessarily invalidate the Varanasis-Blomberg hypothesis, however, It means the Delphinidae/Phocoenidae/Monodontidae merely represent a special case, for there are ely other melon and jaw lipid compositions can successfully function in echolocation, then raises the question of how many dis'e types of lipid composition can be identified mtocete melon and jaw fats. At the present ve can distinguish four groups (Fig. 1). Delphinidae ae head fats contain isovaleric acid, triglycerides and significant amounts (>I 3%) of wax ester, Odonfocefi
Isovoleric
No isovaleric
~d > 3 0 wax
esfers Delphinidae
wax esters
mostly C~o-ci2
Phocoenidae Z~ol~idae Monodontidae
¢q0"C20 oeids
I
Physe~ridae Plafanisfidae
Fig. 1. Comparative lipid patterns in odontocete melon and jaw fats based on analyses of nineteen of the thirtythree living genera. Phocoenidae and Monodontid~[ae head fats also contain isovaleric acid and trigbycerides but little or no (~< 3%) wax ester. Ziphiidae head fa fats possess no isovaleric acid and no distinctive li lipid class composttion; but work now in progresss (Litchfield et al., unpublished) and early distillation lation analyses by Saiki (1953a) indicate that all Zi~iphiidae melon and jaw fats contain large amounts of['C,o--C~fattyacids C not found elsewhere in the Odon toceti. Physeteridae and Platanistidae head lipids possess )ssess no significant amount of isovaleric acid nor av.ly known distinctive lipid class or fatW acid composition.
Since it is the ac erties of odontocete head fats that wou ant in echolocation, compositional data ably be classified on this basis. The v, )und in lipids is a regular function of lax weight (Gouw & Vlugter, 1967) and with the isovaleric acid, C,0-C,s acic chain acid content. On this basis, odol fats can be divided into low (Delphinid ]ae, Monodontidae), intermediate (Zip11 high (Physeteridae, Platanistidae) tool. it categories, each possessmj charact| possessing velocities. Rcpresentative triglyceri (trioctanoin, Cz4), intermediate (trilau intermedi high (triolein, Cs4) carbon numbers n c ~Iz sound waves at 1315, 13: 1357 and 1 ~spectively at 40°C (Gouw & Vlugter, :perimental data are available on sound wax esters, but the same trend tre~ of inct ~y at higher carbon numbers,would be e numbers Can these th three ]ontoceti possessing different acoustical a their head fats be correlatexd with thr ; types of echolocation mechanisms mecln ?( ace is too incomplete to answe wet this qu there is speculative evidence pointing 1 iF n. Norris & Harvey (1972) have ha recently proposed a new t: type of ultrasonic transmission tra for echolocation purposes in Physeter catodon; this involves reflection reflec off the posterior surface of the spermaceti cas~e rather than the unidirectional unidi forward transmissh )n found in dolphins (Norris, 1968; Evans, 1973). The unique convoluti convolutions of the rostrum and mel on in Kogia (Schenkk Schenkkan & Purves, 1973) also suggG:est a distinctive u r nasonic transmission mechani :hanism in that Moreover, Ziphiidae melon tissue tiss structures animal. 1~ and rostr ostrum shapes are known to differ greatly from the corresponding anatomy of the Delphinidae, 1 Phocoenidae, and Monodontidae (N (Norris, 1964; Litchfield & Greenberg, unpublished) blisbed). However, only further study can reveal whether the t~ distinctive head fat compositions of the Ziphiidae, hiidae, Physeteridae and Platanistidae are related to theirr unusual unu cephalic anatomies and their mechanisms of echolocation. echo
Acknowledgements~We extend our th thanks to the following for their help in securing o¢ odontocete fat samples for us in this study: Marineland • ineland of Florida; Dr. E. D. Mitchell, Dr. A. W. Mansfield ant Mr. B. Beck ~field and (Fisheries and Marine Service, Ste. Anne de Bellevue, Canada); Dr. H. Omura and Dr. A. Kawar Kawamura (Whales Research Institute, Tokyo); Dr. P. B. Best (Director of Sea Fisheries, Cape Town); Dr. R. P. van va der Merwe (Fishing Industry Research Institute, Rondebosch, South Africa); Dr. J. G. Mead (Smithsonia fithsonian Institution, University College of Washington, D.C.); Dr. I. Rees (Universi North Wales, Menai Bridge); Dr. H. Olcott Oleo (University of California, Davis); Dr. D. C. Malins and Dr. A. Wohiman (National Marine Fisheries Sela Service, Seattle); Dr. K. S. Norris (University of California, Santa Cruz); Dr. W. J. Houck (California State Univer,sity, Arcata);
D. K. CALDWELL, M. C. CALDWELL, J. eries Service, La x, Canada); and hill, Manitoba). ynthesis of the yses. part by grants (No. GB-30553) ncil.
IFIELD C. (1971) ides and diacyl position ot wax esters, trlglycerldes dlacyl ryl ethers in the jaw and blubber fats of the zon River dolphin (Inia geoffrensis). Lipids 6, 7. N R. G., SIPOS J. C., EATON C. A., HILAMANB . L . ITCHFIELD C. (1973) Molecular species of wax s in the jaw fat of the Atlantic bottlenose dolphin, 'ups truncatus. Lipids 8, 661-667. :AN OIL CHEMISTS' SOCIETY (1967) Official and 7tive Methods, 3rd edn., revised 1971, method -125. American Oil Chemists' Society, Chicago. E. & CANAL H. (1926) Sur l'huile de Mesoplodon ~. C.r. hebd. Seanc. Aead. Sci.,Paris 183,1063-1065. ~RG J. (1972) Pilot whale head oil: lipid analysis ultrasonic studies. G6teborg Congress of the national Society for Fat Research, Abstr. 223. :K T. H., GRINNELL A. D., IKEZONO E., KAMEDA K.ATSUKI Y., NOMOTO M., SATO O., SUGA N. d~ ~,GISAWAK. (1968) Electrophysiological studies of XANAGISAWA •al auditory mechanisms in cetaceans. Z. vergl. central Physiol. 59, 117-156. ~ELL D. K. & CALDWELL M. C. (1971) Sounds CALDWELL produced uced by two rare cetaceans stranded in Florida. Cetology 4, 1-6. EVANS W. E. (1973) Echolocation by marine delphinids one sspecies of fresh-water dolphin. J. Acoust. Soc. and one Am. 54, 191-199. ~N N. K. (1964) Simultaneous determination of FREEMAN trigly¢cerides and cholesterol esters in serum by infrared 5, 236-241. spectophotometry. J. Lipid Res. 5, 1930) Composition of GILL A. H. & TUCKER C. M. (1930) porpoise jaw oil. Oil and Fat Ind. 7, 101-102. 57) Physical properties G o u w T. H. & VLUGTER J. C. (1967) of triglycerides--III. Ultrasonic sound velocity. Fette Seifen Anstrichmitte169, 159-163. 9) Related dietary and HANSEN I. A. & CHEAH C. C. (1969) tissue lipids of the sperm whale. Comp. Biochem. Physiol. 31,757-761. A. (1953) On the study IGARASHIH., ZAMAK. & ISHIGAKIA. of a whale (Ziphius cavirostris, Cuvier) head blubber oil. Bull. Jap. Soc. scient. Fish. 181, 493-496. ). 8r. ACKMAN R . G . JANGAARD P, M., BURGHER R. D. on of the blubber oil (1963) A preliminary investigation hale. J. Fish. Res. Bd from the Atlantic bottlenose whale. Can. 20, 245-247. IA T. KANAZU R., MomI H. & FUKUHARA T., (1969) Studies on est sea area of Kyushu the little toothed whales in the west --XVII. About higher branchedd chain fatty acids in head oil of the little toothed whale--I. Bull. Fac. Fish, Nagasaki Univ. 28, 161-165. Ds J. C. & EATONC.A. LITCI-IFIELDC., ACKMAN R. G., SIPOS (1971a) Isovaleroyl triglycerides from the blubber and Delphinapterus leueas) melon oils of the beluga whale (De Lipids 6, 674-681.
LITCHFIELDC. & Gr I. (19741 Comparative lipid patterns inthe olphins, porpoisesand toothed whales. Cc Physiol. 47B, 401-407, LITCHFIELD C., KAR EENBERG A. J. (1973) Compositional tol melon lipids in the Atlantic bottleno Tursiops truncatus: implications for ecl dar. Biol. 23, 165-169. LITCHFIELD C., KIN/~ :MAN R. G. & EATON C. A. (1971b) C pid patterns in two freshwater dolphi ffrensis and Sotalia fluviatilis. J. Am. C 48, 91A. abolism in fishes--Ill. LOVERNJ. A. (1934) ots. Biochem. J. 28, Selective ,.3t:lectlvt formatio 394-402. KANAZE tty acids in the foetus, MORIIH. nurselin g and adt of dolphin, Stenella attenuata. 7ish. 38, 599-605. attenuat Bull. Jal MORRIS ture of the spermaceti MORRISR. R J. (1973)" seter catodon). Deeporgan of o the spen Sea Res. Res 20, 911-91 NAKAMURA 2) The acidic constiNAKAMUR Y. & T: ¢ dalli True) head oil. tuents cof porpoise ,ric. Chem. Sot ),-645. J. Agric ms of echolocation in NORRIS K. K S. (1964) cetaceans. In coustics (Edited by TAVOLGA -336. Pergamon Press, TAVOLG W. N.), "~ New York. f acoustic mechanisms NORRIS K. S. (1968) ~I ,dontocete ceta( rution and Environment in odon Y University (Edited by DRAKE E. T.), pp. 297-324. Yale Press, New Haven. NORRIS K. K S. (1969) The echolocation of marine maremals. In II The Biology of Marine Mamm, Mammals (Edited by ANDERSEN H. T.), pp. 391-423. Academic Acaderr Press, New ANDERS York. NORRIS K. K S. & HARVEY G. W. (1972) A theory for the function functior of the spermaceti organ of the sperm whale (Physeter catodon L.). In Animal Orientation and Navigation (Edited by GALLER S. R., SCHMIDTI; E,, pp. 397KOENIGK., JACOBSG. J. & BELLEVILLE R. 417. N.A.S.A., Washington, D.C. GARWAL C. V. (1956) PATHAK S. P., SUWAL P. N. & AGARWAI. liver fats. Component acids of suinsh blubber and a Biochem. J. 62, 634--637. on a PERRINW. F. & HtmBS C. L. (1969) Observations Obs( young pygmy killer whale (Eeresa attenuata attenua Gray) from the eastern tropical Pacific Ocean. Trans. Tra San Diego Sue. Nat. Hist. 15, 297-308. RICE D. W. & SCHEFFER V. B. (1968) A 1List of Marine Mammals of the World. Special Scientific Scien Report-Service, Fisheries No. 579, U.S. Fish and Wildlife Wi Washington, D.C. ROBISCH P. A., MALINS D. C., BEST R. (~ & VARANASI U. acoustic (1972) Differences in triacylglycerols from 1 tissues and posterior cranial blubber of o the narwhal (Monodon monoceros). Biochem.J. 130, 33P-34P. 3 SAIKI M. (1953a) Studies on whale,- oil--VII. oil--V Fatty acid composition of bottlenose whale oil--I. On the head and jaw oil. Bull. Jap. Soc. scient. Fish. 18, 1! 497-501. oil--VIII. Fatty acid SAIKIM. (1953b) Studies on whale oiI--VI On the composition of Pacific beaked whale oil--2. o blubber oil. Bull. Jap. Soc. scient. Fish. 19, 1! 809-812. 19731 The SCHENKKANE. J. & PURVESP. E. (1973) Th comparative of the anatomy of the nasal tract and the function f~ ~eteridae (Manmaalia, spermaceti organ in the Physeteridae ferkunde 43, 93-112. Odontoceti). Bijdragen Tot de Dierkunde
~id patterns in odontocete fatty tissues analysis of optiChem. Soc. 42, on the body oils rrue landed in 3f body oils and Soc. scient. Fish.
t components of 7Vhales Res. Inst.
)u~
vtuuu~wz
vii
vt
~Jo.115~,o
d component of Sci. *Jut.
x-~tvv,,x r dolphin. t.tVlVllltt.
Whales Res. Inst. 24, 11%125. gI H. & ITOH S. (1973) Fatty acid components of her oil of the Amazon River dolphin. ScL Rep. les Res. Inst. 25, 293-299. ASI U. & MALINS D. C. (1970a) Unusual wax s from the mandibular canal of the porpoise flops gilli). Biochemistry9, 3629-3631.
VARANASXU. & MAt linked lipids in th (Phocoena phocoen( glycerolipids. Bioc~ VARANASI U. (~ MALl
porpoise (Tursiops, and wax esters o melon) and blubb( 231, 415-418. VARANASIU. & MA characteristic of p structures of diiso 176, 926-928. WEDMm Y., Y LrrcHF~ MITCHELL MITcHEJ E. D. (1 the sition within ss whale. Biochim. bi(
Word Index Key W porpoises; whales.
70b) Ester and ethercanal of a porpoise e of isovaleric acid in 6-4579. [) Unique lipids of the :es in triacyl glycerols ~ndibular canal and ochem, biophys. Acta L972) Triacylglycerols tic tissues: molecular ides. Science, Wash. N R. G., Sreos J. C. & ;neity of lipid comport tissue of the pilot 5, 439-447. ty tissue; dolphins;