On the Giant Octopus (Octopus giganteus) and the Bermuda Blob: Homage to A. E. Verrill

12 Pages·1995·5.9 MB·English

by  S K Pierce

Reference: Dial. Bull 188: 219-230. (April, 1995)
On
the Giant Octopus (Octopus giganteus) and
the Bermuda Blob: Homage to A. E. Verrill
SIDNEY K. PIERCE1 GERALD N. SMITH, JR.2 TIMOTHY K. MAUGEL1
, AND EUGENIE CLARK, ,
1
^Department ofZoology, University ofMaryland, College Park. Maryland20742: and
2Department ofMedicine, Division ofRheumatology, Indiana University
School ofMedicine. Indianapolis. Indiana 46202
"Thesubstancelookslikeblubber, andsmellslikeblubber ofthe sand, haul it up above the high tides (a task that
andit is blubber, nothing more orless." required the efforts of several horses and men), and to
F. A. Lucas. 1897 inform the scientific community ofitsexistence. Because
Webb believed the carcass to be the remains of a huge
Abstract. We have obtained samples oftwo large car- octopus, one of the scientists he contacted was the pre-
casses. One washed up on a beach in St. Augustine, Flor- eminent invertebrate naturalist. Professor A. E. Verrill at
ida, in 1896 and has been occasionally attributed to a Yale(Fig. 1). Based atfirstonly upon aletterfrom Webb,
species ofgigantic octopus (Octopusgiganteus). The other Verrill reported the finding, speculating that it might be
carcasswashedupon Bermuda in 1988 and hasremained aspecimen ofArchiteuthis(Veml\, 1897a). Shortlythere-
unidentified, although its gross morphology, except for a after, having received additional correspondence and
much smaller total mass, was remarkably similar to the photographs from Webb, Verrill concluded that it was a
Florida carcass. We have subjected both samples to elec- "true Octopus, of colossal size . . . one of those upon
tron microscopic and biochemical analyses. Our results which the sperm whale feeds regularly." On the basis of
show that both carcasses are masses ofvirtually pure col- the descriptive and photographic evidence, Verrill pro-
lagen. Furthermore, neither sample has the biochemical posed to name the species Octopus giganteus (Verrill,
characteristics ofinvertebrate collagen, nor the collagen 1897b). However, almost immediately, Verrill changed
fiber arrangement of octopus mantle. Instead, they are his mind. Based upon more photographs, measurements,
large pieces ofvertebrate skin, the Bermuda sample from and descriptions ofthe carcass after it had been entirely
a poikilotherm and the Florida sample from a huge ho- unearthed from the beach sand, along with several for-
miotherm. We conclude that there is no evidence to sup- malin-preserved pieces ofthe tissue, Verrill retracted his
port the existence ofOctopus giganteus. rapidly drawn, initial conclusions, writing that "thecrea-
turecould not have been an Octopus" (Verrill, 1897c, d).
Instead, Verrill, togetherwithsome otherbiologistsofthe
Introduction time, came to the conclusion that the carcass was from a
The first evidence that seemed to document the exis- large vertebrate, most likelyawhale(Lucas. 1897; Verrill,
tence ofa species ofgigantic octopus washed ashore on 1897d, e). However, otherbiologists, notably Dr. William
ayenarocoefan1i8c96b.eaTchhe asto-Scta.llAeudgucsartcianses,wFalsortidaak,enlaitne cihnartghee MH.usDealulm, otfheNnattuhrealCuHrisattoorry,osftilMloflalvuosrkesd Vaetrrtihlel'sNaotriigoinnaall
species diagnosis according to correspondence between
by a local physician. Dr. DeWitt Webb, who was also him, Webb, and Verrill (archived at the Smithsonian In-
president ofthe St. Augustine Scientific Society (Wood,
1971). Webb set about to photograph the body, dig itout stiAtulttihono,ugWhasVheirnrigltlond.isDaCv)o.wed his species description of
Octopus giganteus several times, the carcass was never
Received 31 October 1994;accepted 25January 1995. properly identified. The matter rested quietly for 70 years.
219
220 S K. PIERCE ET AL
ofcellular structure found in it by Gennaro (1971) (but
not the lack ofsuch structure in the "contemporary" tis-
sues) and the persistence of the carcass as it lay on the
beach. (According to the correspondence between Webb
and Dall, the carcass was still on the beach on March 17,
1897, almost 4 monthsafterthe initial discovery.) Mackal
( 1986) concluded that the amino acid data together with
some inconclusive Cu and Fe measurements "support the
original identification ofthe tissue and carcass by A. E.
Verrill asanexceptionally largecephalopod, probablyoc-
topus, not referable to any known species," in spite of
both VerriU's change ofmind and the complete lack ofa
suitable test oftaxonomic relationships in Mackal'sdata.
In the end, while theexistenceofthe St. Augustine carcass
is well documented and the discovery often cited [most
recently in the popular press (Ellis, 1994) and in a bio-
logical science text book (Milne, 1995)], there is no un-
equivocalevidenceatallthat itbelongedtoagiantoctopus
or, indeed, to any particular species.
During the summer of 1988 another carcass washed
Figure 1. A. E. Verrill.Courtesyofthe Marine Biological Laboratory into a lagoon on the island of Bermuda. This unrecog-
Archives. nizable carcass 2.50 X 1.25 X 0.30 m), immediately
(
labeled the "Bermuda Blob" by the popular press, was
and then a report appeared stating "it can be safely said lpohcoatlodgirvaeprhaenddafnidsshearmmpalnedwhbyoToeftdednywTourckkserw,itahrsecnieonwtnisetds
that the gigantic mass of tissue that washed up on the on Bermuda. Whileconsiderably smallerthan theSt. Au-
beach at St. Augustine in 1896 was the remains of an
o1c9t71o)p.usT.ha.t.r2ep0o0rtfeceton.ti. n.u"ebdettowedeetnaitlentthaeclcehrtiopnso(lWoogyodo,f gscursitpitnieondiexsaccotvleyr,y,ontahegrBosesrlmeuvedla. "cNaorcabsosnefsitoVrehrrairldl'psardtes.-
theevents summarized above and also indicated that siz- t. o.ug.hInasntdeadelaosftibce,inagndmucsocmuploarse.d. m.a[itnhleytiosfsumesu]cahreifnitremr,-
achbeollmedpaaptriietcsheeosnSomofifttthhhseeonctiaisrascnuaesIsnfsrthoiatmduttibhoeene.SnAmipctruherssseoornrviyeadhniaswtniodtlhogwtiehcraaelt ln.ae.cc.etdi.vfeSibtoeirmssseuaen.lda[rglTeahregiertribesugsunuldealries]scadonifaffltisocuuplgtehrtfmoiebcarutotueso,trhwetheia[trteisacspuoaenr]-t.
of "contemporary" squid and octopus was carried out. These may havecontained blood vesselsoriginally. From
Although the specific squid and octopus tissuesthat were theinnersurface ofsomeofthepieceslargecordsofelastic
compared to the Florida carcass were not reported, no fibers proceeded inward" (Verrill, 1897c). Gennaro's ad-
cellular structure was found in any ofthe tissues. Instead,
ditional description ofthe St. Augustine tissue also fit the
arecvoenanleedctwiivtehtipsosluaerifizbeedr lniegthtw.orTkheincaolnlctlhurseieontsissrueeascwhaesd nBeecrtmivuedtaiscsaurecwasassesxoacttoluyg.h"tWhhatititedausllseodafpou.r.bl.atdheesc.o.n-.
were that none ofthe samples looked mammalian, that the same homogenous, tough, white, fibrous texture
the St. Augustine tissue looked much more like the oc- [throughout]" (Gennaro, 1971).
topus fiber network than that ofthe squid, and therefore, We have been able to obtain small pieces ofboth the
"the St. Augustine sea monster was in fact an octopus" St. Augustine and Bermuda carcasses. We have subjected
(Gennaro, 1971). Because the report ofthese histological both to electron microscopic examination as well as bio-
studies was written for a general, rather than scientific chemical analyses to test the similarity between the two
audience, it lacked a rigorous description ofprotocol and tissue masses and to determine their taxonomic origin.
observations. In addition, we have carried out light and electron mi-
The matter rested again, this time for another decade croscopicexaminationsofoctopus(Bathypolypusarcticus)
and a half, until the appearance of a report about the mantle tissueand humpback whale(Megaptera novaean-
amino acid composition ofan acid hydrolysate ofthe St. gclue) blubber.
Augustine tissue, by now almost 100 years old. Although
Materials and Methods
neither hydroxylysine nor hydroxyproline concentrations
mn
were determined, the amino acid composition suggested Elect microscopy
that the St. Augustine tissue was likely to have been a Bermuda andSt. AugustineCarcasses. Specimens from
huge massofcollagen (Mackal, 1986),explainingthelack both the Bermuda and St. Augustine tissue masses were
THE GIANT OCTOPUS AND THE BERMUDA BLOB 221
SillSK'.fcAi'S "^jSfa&jSfeVj-kSW&SS',.:
Figure 2. Low magnification transmission electron micrographsofsectionsofthe Bermuda(A)and St.
Augustine(B)carcasses.Thecollagenfibersofbothtissuesruninlayersthatareperpendiculartoeachother.
Within each layerthe fibersappearto beorganized in bundles(seethe upperhalfofA). Thistypeoffiber
organization is typical ofskin collagen (see Discussion). Other than the fibers, no othercellular elements
werefound.Bacteriaandbacterialspores(arrows)werescatteredthroughoutthefiberlayersinbothsamples.
prepared for electron microscopic examination. Although followed by 0.2% lead citrate for 1.5 min (Venable and
theexactcomposition oftheoriginal preservation medium Coggeshall, 1965). The stained sections were examined
EM
was unknown for either tissue, the distinct odor of for- with a transmission electron microscope (Zeiss 10
malin was obvious in both. Indeed, Webb, in his corre- CA) at 80 kV.
spondence with Dall, indicated that he had put several Octopus nuinlle. The specimen ofB. arcticus (USNM
pieces ofthe body in formalin before sending them offto catalogue #884184) that provided a mantle tissue sample
both Dall and Verrill (cited in Wood, 1971). We cut sev- hadbeencollectedbytrawlofftheNewJerseycoastduring
eral pieces ( 1 mm1) from each ofthe original tissue sam- a 1981 cruiseofthe R/V DelawareIIand hadbeen placed
ples and placed them directlyMinto 2.5% glutaraldehyde/ immediately into formalin upon its capture. At some
2% paraformaldehyde in 0.1 cacodylate buffer con- point, the octopus was transferred into isopropyl alcohol
M
tainingO.3 sucrose(~1 100 Mmosm) (Bermuda sample) and had been stored in that solution until we were given
or 2% glutarMaldehyde in 0.15 cacodylate buffer con- accesstoit. Wecutasectionofthemantleofftheoctopus,
taining0.58 sucrose (~ 1 100 mosm) (Florida sample). rehydrated it and placed it into 2% glutaraldehyde. The
The tissue samples remained in the fixative for several tissue was then prepared for electron microscopy, as de-
days. After fixation, the tissue pieces were post-fixed in scribedabovefortheFloridasample. Forlightmicroscopy,
2% OsO4 in the cacodylate-sucrose buffer, treated en bloc thick sections were cut from the same specimen, stained
with 2% aqueous uranyl acetate, dehydrated in an ethanol with Richardson'sstain (a mixture ofmethylene blueand
series (35-100%), and embedded in epoxy resin (Spurr's Azure II). and viewed with bright-field optics (Zeiss. Pho-
medium). Thinsectionsshowingasilverinterferencecolor tomicroscope II).
were cut with a diamond knife on an ultramicrotome Whaleblubber. Wecutasmall sample ofblubberfrom
(Reichert, Ultracut E). The sections were mounted on a much larger piece that came from a male humpback
coppergridsand stained with 2% uranyl acetate for 5 min. whale that had died at sea and washed onto a beach in
222 S. K. PIERCE /:'/ I/
Figure 3. Higher magnification transmission electron micrographs showing the periodicity along the
fibers n!"the St. Augustine collagen (A), rat tail tendon collagen (B). and the Bermuda collagen (C). all at
thesame magnification. ThefibersoftheSt. Augustineand Bermuda samplesarethinnerthanthosein the
rattail tendon acharacteristicofskincollagen. Thesomewhat indistinct bandingpattern ofthe Bermuda
fibers is hkcl\ due to the poor original fixation. The dense deposits in this sample also derive from the
original fi\ati\e solution.
THE GIANT OCTOPUS AND THE BERMUDA BLOB 223
Figure 4. Light micrographs ofcross sections taken through the width ofthe mantle ofBathypolypus
arcliais. The epidermis, which consists ofan epithelium underlain with a thin layer ofdispersed collagen
fibers, hasbeen removed. (Micrograph A) The mantle consists oftwo primary layersofmuscle bundlesof
about equal width. These layersare separated by a spacethat containsblood vessels (oval structure in the
center)(magnification = 62X). (Micrograph B)Theoutersurfaceofthemantleconsistsofsmallbundlesof
longitudinal muscles(L)coveredbyaverythinlayerofcollagen(arrowheadC). Underlyingthelongitudinal
musclesaremusclebundlesthatalsorunlongitudinally,butcontainingindividualfiberswithineachbundle
thatare not parallel to each other(M). Interspersed between these deeperbundles run thin radial muscles
(R)which span the width ofthe mantle (see micrograph A. also) (magnification = 390X). (Micrograph C)
Belowthebloodvessellayerrunadditionalmusclebundlescontainingfiberswithawidearrayoforientations
(M). Small bundles ofcollagen (arrowheads C) containing fibers that run parallel to each other occur oc-
casionally between the muscle bundles. The hollow, tubularstructure ofthe muscle cells isevident in this
section (magnification = 390>). (Micrograph D) The innersurface ofthe mantle isalso covered byathin
layerofcollagen(arrowheadC)thatextendsupwardbetween(arrowC)adjacentbundlesofthelongitudinal
muscles(L). Although notshown in thissection, the radial musclesinsert between the longitudinal muscle
bundles(see micrograph A). The longitudinal musclebundlesare much largeron thisaspect ofthe mantle
than thoseon theoutersurface(comparewith micrograph B). Immediately abovethelongitudinal muscles
runsathin bundleofcircularly oriented muscle fibers(magnification = 390X).
Virginia Beach County, Virginia, in October 1992. Ac- cayed. The original piece ofthe blubber, still attached to
cording to the Virginia Museum ofScience collection re- the epidermis, had been preserved in formalin and de-
cord, this carcass was 906 cm long (about 200 cm longer posited at the Smithsonian (catalogue #VMSM 921025).
than the St. Augustine carcass) and only moderately de- Oursamplewastransferred into1%glutaraldehyde, small
ff ' : , .....-. . ..
'".-^^^'f*' ":>...:''.. ''y^-'A
I
.;;;,
..
Figure 5. Electron micrographs ofBathypolypus arcticus mantle. (Micrograph A) This cross section
shows the arrangement ofthe contractile proteins in bundles that radiate from the hollow center ofeach
tubular musclecell (M). Adjacent muscle fibersare separated by occasional bundles ofcollagen fibers(C).
(Micrograph B)The fiberswithin thecollagenbundles(C)alwaysrun parallel toeachother. Adjacentlayers
ofperpendicularly runningcollagen fiberswere neverseen. (MicrographC)Thecontractile proteinswithin
224
THE GIANT OCTOPUS AND THE BERMUDA BLOB 225
pieces were cut from immediately under the epidermis, droxylated (hydroxyproline, threonine, serine, tyrosine.
the middle, and the inner regions ofthe blubber, and all and hydroxylysine), and polar (aspartate, glutamate, hy-
wereprepared forelectron microscopyasdescribedabove droxylysine, lysine, histidine, ornithine, and arginine)
for the Florida sample. amino acid residuesper 1000in thecollagen hydrolysate.
Rat tail tendon and skin. Because our initial micro- Eachsumwasdividedbythegrandsum ofthethreeamino
scopic examination ofsections from the carcasses found acid groups and then multiplied by 1000 to yield a set of
fibers that resembled collagen, we proceeded to measure three of Matsumura's R values. A vector R is then plot-
the periodicity ofthebandingpattern ofthe fiberstocon- ted on a triangular coordinate graph using the R values
firm that identification. We used collagen fibers from rat (Rhydrophobicj ^hydroxyiated- and Rpo\-dr) as coordinates (Mat-
tail tendon as an internal standard for these measure- sumura, 1972).
ments. A piece oftail was obtained from a white rat that
had been decapitated and immediately frozen for other Results
experimental purposes. The tail was thawed, skinned, and
mm
the tail tendon removed. Pieces (1 3) were cut from The electron microscopy and the amino acid analysis
both theMskin and tendon and fixed in 2% glutaraldehyde indicate that both the Bermuda and St. Augustine car-
in 0.12 phosphate buffer (pH 7.4). Following initial casses are made up almost exclusively ofcollagen fibers.
fixation, thetissue pieceswerewashed in theabovebuffer The pieces oftissue from the Bermuda and St. Augus-
and then postfixed in 2% OsO4 in the phosphate buffer. tine carcasses contain layers of fibers showing banding
Subsequent preparative steps were the same as described patternsthatarecharacteristic ofcollagen, a fewscattered
above for the Florida sample. bacteria and bacterial spores, and no othercellularstruc-
tures (Fig. 2). In both specimens, adjacent layers of the
Amino acidanalysis collagen fibers run perpendiculartoeach other. Although
the fixation ofthe Bermuda sample is not verygood, lon-
In the case ofboth samples, the small piece oftissue gitudinal sectionsofthetwospecimensappearverysimilar
that remained following the microscopy (256 mg, Ber- at low, and even at high, magnifications. Certainly, the
muda, 216mg, St. Augustine) was soaked in several bandingperiodicityalongthefibersissimilarbothtoeach
changesofartificialseawater(940 mosm)overnightat4C otherandtorattail tendon collagen (Fig. 3). We measured
towash outthe preservativesolution. Thetissuewasthen thedistancebetween the majorperiodsalongseveral fibers
mm N
cut into pieces (2 3) and hydrolyzed for 24 h in 6 in several sections at both low and high magnification.
HC1 at 100C. Both samples dissolved within 15 min of The averages were 54.3 nm (2.72 SD, n = 150 mea-
being placed into the acid. The hydrolysate was neutral- surements) for the St. Augustine collagen and 57.9 nm
ized with NaOH and the amino acids extracted with an (5.37 SD, ;; = 154 measurements) for the Bermuda fi-
equal volume of95% ethanol. Theextractwascentrifuged bers. Although these values are slightly less than the usu-
at 20,000 Xg(4C), andthesupernatantwasfreeze-dried ally published periodicity for collagen banding (60 nm),
overnight. The residue was dissolved in 0.2 N lithium control samples ofboth rat tail tendon (Fig. 3B) and rat
citrate buffer (pH 2.2), and the amino acid composition skin collagen yielded a banding periodicity of 55.7 nm
ofthis last solution determined with an automatic amino (5.6 SD) with our protocol. The diameter ofthe indi-
acid analyzerwith ninhydrin detection (Beckman. System vidual fibers was also determined from micrographs of
Gold). Amino acidconcentrations werecalculated bythe bothcrosssectionsand longitudinal sections. Theaverage
SystemGoldsoftwarewith norleucineasan internalstan- for the St. Augustine collagen was 109 nm (25.2 SD, n
dard, and then converted to residues/1000 residues for = 68), 156 nm (34.7 SD, ;; = 106) forthe Bermuda col-
each individual amino acid. These data were graphically lagen, and 173 nm (82.0 SD, n = 31) forrat tail tendon
comparedtotheamino acidcompositions ofthecollagens collagen.
of97 speciesfrom diverse phyla, accordingto the protocol Microscopic examination of Bathypolypm arcticus
described by Matsumura (1972). mantle revealed a structure that is dramatically different
Briefly, the Matsumura protocol consistsofcalculating from that ofthe two carcasses. In particular, the massive,
the sum ofeach ofthe hydrophobic (valine, methionine, perpendicular collagen fiber arrangement characteristic
leucine, isoleucine, tyrosine, and phenylalanine), hy- ofthe two carcasses is completely absent in the octopus
the sarcomeresofthe mantle muscle cellsare not in register. (Micrograph D). The radial muscles(R) that
span thewidthofthe mantlebetween therestofthe mantlemusculature(M)areattached tothe innerand
outer layerofcollagen (C) byjunctional complexes (J). The dorsal surface ofthe mantle liesto the left of
thismicrograph.
V
<
*j
W
V
5=5=3,^^ ';> 'V; :,: .
;',''; ., '*;'
5-^W^.^IS^S
ir.-V'" .v:&i''?'iv jL-ii.i^ "s?f -.,''4:'V~
Figure6. Electron micrographsofhlubberfrom McKtiplcru novaeangelae. (Micrograph A taken near
theepidermis) Large bundlesofperpendicularlyoriented collagen libersare interspersed withcells(mostof
which appeartobeh'hroblasts), lipiddeposits(L),andoccasionalelastin fibers(E).(Micrograph B)Collagen
226
THE GIANT OCTOPUS AND THE BERMUDA BLOB 227
mantle. Instead, the bulk ofthe mantle is composed of contractile filaments within each sarcomere are not
muscle. Small amounts of collagen are located in thin striated (Fig. 5C). Thick filaments, reminiscent of para-
sheets, one between the epithelial cell layer ofthe epider- myosin, surrounded bythin filamentsareevident incross
misand theoutersurfaceofthe mantle, the othercovering sections of the muscle cells. Squid mantle muscle cells
the innersurface ofthe mantle. In addition, collagen also have a similar fine structure (Ward and Wainwright,
occurs in an internal network ofsmall bundlesofparallel 1972).
fibers running between the muscle bundles (Fig. 4). The Microscopic examination of the blubber from the
banding periodicity alongthe octopus collagen fiberswas humpback whale revealed a massive matrix ofcollagen
46.6 nm (8.6 SD, n = 90), 15% smaller than the peri- fiberspresent throughout theentirethicknessofthetissue
odicity ofthe St. Augustine collagen. (Fig. 6). Fat deposits and poorly fixed cellular structure
The muscle fibers that make up the bulk ofthe mantle were evident between layers ofcollagen fibers in the sec-
arearranged in several layers (Fig. 4). Immediately under tionsoftheblubbertaken both close totheepidermisand
a thin external collagen layer is a layer ofparallel muscle from the medial region (Fig. 6A, C). Neither fat nor any
fibersthat runslongitudinally from themantleedge. These remnantofcellularstructurewasfound ineithertheFlor-
fibers are separated laterally from each other by thin idaorBermuda carcass. The sectionstaken from the inner
downward extensions ofthe coveringcollagen sheet (Fig. aspect ofthe blubber layercontained only collagen fibers
4B). A similar muscle layer, although consisting oflarger in very large bundles, interspersed with occasional larger
diameter fibers, is located immediately inside the collagen fibers that appeared to be elastin (Fig. 6D). At all levels
layerthat formstheinnersurface ofthe mantle (Fig. 4D). ofsection, the collagen fiber arrangement ofthe blubber
The center ofthe mantle is occupied by a space that con- was exactly that ofthe Florida and Bermuda carcasses,
tains blood vessels (Fig. 4A). Ventrally between the namely, bundles arranged in layers runningperpendicular
blood vessel space and the inner, longitudinal muscle toadjacent layers(Fig. 6A, B,C). Thebandingperiodicity
layer lie several layers ofmuscle fibers that run at a va- along the whale collagen fibers was 54.6 nm (5.1 SD, /;
riety ofangles obliqueto thecross-section (Fig. 4C). Dor- = 83), essentially identical to the banding ofthe St. Au-
sally, between the blood vessel space and the outer lon- gustine collagen.
gitudinal muscle layer, the muscle fibers generally run in The amino acid analyses of the tissues from the two
thesamedirectionasthefibersoftheoutermusclebundles carcasses were also suggestive of collagen. Glycine ac-
(Fig. 4B). Lastly, the outer and inner collagen sheets are counts for about one third ofthe amino acid residues in
occasionally connected toeach otherbythin, radial mus- both tissues, and both hydroxylysine and hydroxyproline
clebundlesthat traversethewidth ofthe mantle, perpen- were present as well; these features are virtually diagnostic
dicularto the rest ofthe musculature and attached to the ofcollagen (Table I). However, the amino acid compo-
collagen sheets by ajunctional complex (Figs. 4A, B, C; sitions ofthe hydrolysates ofthe two carcasses are quite
5D). All ofthis structure is supported by occasional thin different from each other. In particular, the St. Augustine
bundles of parallel-running collagen fibers (Fig. 5A, B). carcass is very rich in proline (169 residues/1000) and
Altogether, there is nothing in the octopus mantle mor- quite low in lysine (0.4 residues/1000), in comparison to
phology that resembles anything in the two relics. both the Bermuda collagen (88 residues/1000 and 10 res-
The mantle muscle cells are basically hollow tubes, ta- idues/1000, respectively) and skin collagens from several
pered at each end. The contractile proteins are arranged otherspecies(Table I). Ofcourse, the unusually lowlysine
in bundlesthat arerectangularin cross-section and radiate values in the St. Augustine sample may be an artifact of
out from the hollow center ofthe cell like tightly packed a century in formalin. Only whale skin collagen (species
spokes (Fig. 5A). The nucleus and mitochondria are lo- not reported in Eastoe, 1955) has proline residues/1000
cated in the center ofthe cell. The arrangement of sar- that are anywhere near those from the St. Augustine col-
comeres in the octopus mantle muscle gives the appear- lagen (Table I). In addition, the amino acid composition
ance ofoblique striations in occasional sections, but the ofthe St. Augustine collagen is very different from that
fibersarrangedinbundlesrunningperpendiculartoeachotherareeverywherethroughouttheentirethickness
ofthe blubber. The size ofthe bundles varies, depending upon location within the width ofthe blubber.
(MicrographC from the middle oftheblubber layer). The lipid deposits(L) in thisregion ofthe blubber
werelargerandmorefrequentthantheotherareasexamined.Theperpendiculararrangementofthecollagen
bundlessurroundingthe fatdepositswasstillevident(lowerrighthandcornerand upperleft handcorner).
(Micrograph D taken from the inneraspectoftheblubberlayer)Thecollagen bundleswereverylarge in
thisregion.Theexpanseofcollagen fibersshown hereareall incrosssection,andwerebounded byequally
largeexpansesofperpendicularly runningcollagen fibers. Veryfewcellsorlipiddepositswereencountered
in thisregion oftheblubber, althoughelastin fibers(E)werequitecommon.
228 S. K. PIERCE ET AL.
Table I
Comparativeaminoacidcompositions ofskincollagensofseveralspeciesandthe BermudaamiSi. Augustinecarcasses
(valuesareaminoacidresidues/1000 residues)
Ammo
acid