On the Pigmentation of a Clypeastroid, Mellita sesquiperforatus Leske

W. J. Crozier

The American Naturalist, Vol. 52, No. 622/623. (Oct. - Nov., 1918), pp. 552-556.

Stable URL:
http://links.jstor.org/sici?sici=0003-0147%28191810%2F11%2952%3A622%2F623%3C552%3AOTPOAC%3E2.0.CO%3B2-M

The American Naturalist is currently published by The University of Chicago Press.

Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at
http://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained
prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in
the JSTOR archive only for your personal, non-commercial use.

Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at
http://www.jstor.org/journals/ucpress.html.

Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed
page of such transmission.

The JSTOR Archive is a trusted digital repository providing for long-term preservation and access to leading academic
journals and scholarly literature from around the world. The Archive is supported by libraries, scholarly societies, publishers,
and foundations. It is an initiative of JSTOR, a not-for-profit organization with a mission to help the scholarly community take
advantage of advances in technology. For more information regarding JSTOR, please contact support@jstor.org.

http://www.jstor.org
Fri Dec 14 15:33:16 2007
552 TIIE A X E R I C A S YATCRALIIST [T'oI,. LII

that their notes were delivered inclependentlg of the rhythlii of

this particular group. From observations of tlie striclnlations
of other groups i11 this sanie colony, I an1 of the opinion that it
is not unusual for these locusts to develop a rhythinic synchro-
llislil ill sliiall groups.
I t ~voulclbe interestilig to k11o~vwliy sonle species of locusts
ancl crickets possessing the iiitermitteiit habit of striclulation
tend to develop a more or less perfect r h y t h n ~ i csynchronisln
~ v l ~ i others
le do not. Although this is true of the two cricliets,
CEccriztl~uswivezrs and C y r t o z i p h a c o l z ~ ~ i ~ b i aIn ahave
, been unable
to note any syachronisni in the chirpings of the conlinon arboreal
cricli-et, Orochnris saltator. Although large colonies of these
crickets liiay often be heard in striclulation, each individual
appears to striclnlate in its orrn leisurely liianlier independently
of its fellows.
H. A. ALLARD
TVASHII~GTOK, D. C.

ON T H E P I G N E N T A T I O N O F A CLYPEASTROID,
MELLITA SESQUIPERFORATUS L E S K E 1
THE colnmoll clypeastroids, E c l ~ i ~ ~ c i ~ ) ' c iand
c h ~llIcllitn,
~ i ~ ~ s nrllen
aclnlt, are characteristically of a broxvn or (in the former spe-
cies) redclish-bro~vncolor. This seenis to be general throughout
the group. Taxonomic lists contain, ho~vever,numerous refer-
ences to a greeaisl~ coloratioll of the test of these animals.
W11en preserved in alcohol, or when dried, either after fixation
in alcohol or after killing with fresh water, these sand dollars
usually assume, for a time a t least, a somewhat greenish color.
Clarli- (1899, p. 118) says lhat speciinens of ,llellita sesqz~iper-
foratz~s Leske (=sexforis A. Ag.) collected a t Jamaica were
delicate olive green [when alive, I infer, thong11 with doubt].
He also gives the coloratioll of specimens of this species obtained
in Porto Rican waters as "usually light olive green (rarely
brown) whell alive." A t Berinnda living incliviclnals of this
species are, he says, invariably browa, with no llint of green
about them, either 011 the esterilal surface or in the viscera.
TT711en killed in alcohol, however, they become green, and greea-
ish piglneilt is dissolved by the Anid. This is also true of E c h i -
?zarachnizis parnza (Clark, 1904, p. 564; Coe, 1912, p. 111).
No~ir, examination shomrs that there is a t the bottom of this
1 Co~ltributiollsfrom the Bermuda Biological Station for Research! No. 90.
Nos. 622-6231 X H O R T E R A R T I C L E S AAIrDDISCUSLSIO,\~ 553
matter-in descriptive lists sonlemhat confnsing-a rather in-
teresting point, which it is the purpose of this ~ l o t eto elucidate.
~llellitn,a dult, is at Bermucla undoubtedly b r o ~ v n large
; speci-
nlens (9.0-11.5 em. in transverse diameter, usually 9.5 em.),
~vhichI have from time to time collected by dredging upon
grass-free bottonls of fine sand or mud a t Flatt's Inlet, Spanish
Point, TTVORock Passage, and other localities, are uilifornlly
bromn ~1po11both aboral and oral s-t~rfaces,although the different
iilclividuals vary somewhat as to shade. Their general hue
liarmonizes well with that of the substratum. I t is improbable
that light has had a direct effect in producing pigmentation,
since the oral surface, neyer turned toward the light, is at least
as densely pigmented as the aboral one and is freclnently ( i n
larger specimens) clarker. Youilg individuals i n an active,
healthy state mere gotten in associatioil with aclnlts during the
antninn months. U p to 5 em, diameter, i11 one case 8 em., they
were found, with one exception in about 30, to exhibit no bromn-
ish coloration wliatever; they were, on the contrary, g ~ i r ewhite,
and translucent, the yellowish stonlach being easily made out
through the test. These individuals were usually 3.5 to 4.0 em.
in transverse diameter. The one exceptiollal specimen, 2 em.
in diameter, mas ~ u l u s n a lbecause it was of a light coffee-brown
shade.
TTT1len placed in alcohol, or in fresh water, these young white
Jlellitas became bright green; i n s~ulliglltthe green on aleoliolic
specinlens cjuickly disappears. C l a ~ l i(1901, p. 254) notes that
sonie young specimens of N . pentapora exanlined by hi111 were
green [in alcohol ?I ].

TTrhen kept in aclunria for several days the imall uliite "scw-
plates" developed, in most cases. local indicatiolls of green pig-
ment, altlioagh the animals were still quite active. Tliis was
also true of the large b r o ~ v nindividuals. I t mas noticed that
in cases where a large brosvn Jlellita had been damaged by the
cutting eclge of the dredge, a green coloration was present along
the ~ v o u n dwhen the haul was brought to the surface. Other
specimens, apparently uninjured, sonietilnes developed an olive-
green color on the oral surface within half an hour after being
transferred from the dredge to a tub of sea water.
Thus the green coloration of illellita is associated svith a con-
dition of injury or cleath. I t is possible that the green Inaterial
may have no connection with the substance responsible for the
general brown integumentary coloration of adults.
564 TITE A~~~~~~~~~ ,I~ATCR~ILI~T [TJOL.
LZI

The green pigilient is readily extracted ~vitlifresli ~vater. It

is not chlorophyll. Tylien an ailinla1 is allosvecl to clie in fresli
water, tlie integument and the ordiilarily white internal p a r t i
of the slceleton of a il1ellitn become bright green. Putrefactive
changes clecolorize the green extract, anel the color call not be
restored by alkali, or ~ y i t hH,02. Green extracts are also cle-
con~posecl,irreversibly, by boiling.
The green color is not seen in faintly acid fresh-~vaterextracts
and the Xcllitc~1+emaiiis bro~vn. If such an extract is made
alkaline, tlie green color prolnptly appears in tlie extract. The
substance responsible for the green llne in cleat1 or injured parts
of llIell~tc~ is in fact a very goocl indicator. It is colorless in
acid, vivid green in alkalilie solutions; this color change may be
lnally times. The (turning-point" of the illdieator
(

is at an acidity of p,=7.6-7.8-in a solutioll more alkaline

than nentrality, but well on the acid side of tlle reaction of sea
water ( ' p ~ - - 8 . 1 t ) . This greeiling material seems to be pres-
ent throughout the body of Lllellitn, as freshly secured bits of the
(white) internal skeleton t u r n bright green in alkali. The felt7
available references to the coloration of clypeastroids indicate
that the allrali-greening substance regularly occurs in Clypenster
and in other genera of this group.
The ovaries of are heavily pigillented by
ccsqz~iperforail~s
a substance of the "antec1onin"-"echinocliroii~e" group. The
mature egg itself is light brownish yellow, heavily yollrecl, and
apparently larger tlian any echinoid egg that has been described.
I t measures abont 0.26 mm, in diameter, and is thus abont twice
the size of the egg of 11. testz~di)znict (=pe)ltaporn), which
ineasnres 0.11 inm., though not so large as that of the brittle-
star Ophioder?ilu, 0.30 111111. in diameter (Grave, 1916, p. 439).
The ovarian egg of If. cecqzlipcrforcttus is surrounded by a
gelatinous envelope, the wliole being 0.35 t n1m. in diameter.
This envelope bears numerous evenly scattered clulllps of pre-
cipitated reddish-purple pigment, the ovarian stronla being
densely crowded with similar " chron~atophores." I11 Echi-
?~nrachnil!sthe LLcl~romatopl~ores" of the egg-envelope are red
rather than purple. The purple pignzent of the J1ellitn ovaries
exhibits the acid-alkali color changes and the absorption spec-
trum of the " echinochrome" pignlents found ia sea-urchins,
holothurians, crinoids, and even ill star-fishes; a dilute extract
of a fenlale Xellitn may therefore be prepared which changes
Nos. 622-6231 XHORZ'ER .IRZ'IC LES ,LAYDDISC'chSIO.'lr 555
from bright green to reddish purple at a hydrogen-ion concen-
tration of about 7.3, the L'echinocl~romc"not beco~lliligorange
until a nluch greater acidity is reached. Thii pigment becomes
blue at a11 allialinity of about p,,= 8.2; in the ovary it is recl-
dish purple, but it is not in solution.
The interest of these facts lies not so much in their supplying
some adclitiollal i~lstallcesof tlie elaboration of similar (or even !
identical) snbstances-cons1~ic11o11sly pigments-by animals re-
latecl in descent, but in the eviclellce which is aflorded regard-
ing the reaction of intracellalar fluids. Xumerous cases illns-
trating the former poiat are available fro111 alllolig echiaoclerlns,
molluscs, tunicates, a n ~ so l forth, and tliese cases have a certain
impol.tance for the general theory of animal coloratioa. But I
am here chiefly concerllecl to point out that, if the alkali-greeaing
substance present in J1. sesqziiperforatz~s is closely similar to
that produced in the tissues of other clypeastroicls, nllequivocal
statements as to the occurrence of greellish hues in living "sand
clollars" may contain a sugge~tioaas to a possible Illode of origin
for certain 1~11o~~711 geographical color diflerences in echi~loderlll
species. I have not been able to find such references, but tlie
point is worthy of further study. The evidence afforded by
iatracellular substa~lces capable of behaving as indicators of
acidity shows plainly that the tissues of nlarille animals are
nlucli nlore acid (less allialine) than sea water. I n JI. sesqlii-
p c r f o r n t l ~ s the green color is produced when the tissue fluids
become more allialine than they customarily a r e ; thus tlie in-
tegument, svlien injured or Billed, becomes permeable to sea
water and assmnes a green hue. Healthy indivicluals for rs-
perinzelltal work may, incidentally, be selected (at Bermuda)
by the absence of green areas upon the test; the readiness svitli
1~-11ichgreening is induced ilidicates the degree of care which
must be enlployed in haadling some marine animals. If by
some means, for example, by higher temperature, the tissues of
a Mellitn population i a a ~~~ar111er sea svcre constantly main-
tainecl at a higher alkalinity than those at Bermuda, they might
normally appear somesvhat greeliish in color. The normal varia-
tions in the coloratioll of Chro?)zoclorisxebrcr, a nudibranch con-
taiaing a11 illdieator favorable for such ubservatioas, strongly
suggest that such differences in the reaction of intracellular
fluids (not necessarily of the protoplasm) are entirely possible
(Crozier, 1916). Whether or not conlparable changes may be
 TEE AJ1ERICBAL7S,lTURdLIS'T

iiiduced as a regular thing i a different oceanographic regions

call not as yet be stated.

REI'EBESCES
Coe, TY. R.
1912. Echiilodernis of Connecticut. S t a t e Geol. a n d N a t . IIist.
Snrv., Bull. 19, 152 pp., ills.
Clark, EI. L.
1599. F ~ l r t l i e i K o t e s on the Echinodclnis of B e ~ m n d a . Aiut. AT. Y.
Acnd. Xci., Tol. 12, pp. l l i - 1 3 8 .
1901. The Echinoclerm~ of P o r t o Rico. Bull. U. S . F i s h Conlm, f o r
1900, Tol. 2, pp. 231-263.
1904. The Echinoclernls of t h e TToods Hole Region. Ibiil., f o r 1902,
pp. 545-576.
Ckozier, TV. J.
1 9 1 6 ~ . Some Inclicntois froin l n i n l a l Tiwnes. Jour. Biol. Che~n.,
Yol. 21, pp. 413-445.
19168. Cell Pelletlation by dcicls. I T . F u r t h e r O b - e r ~ a t i o n s 011 the
Blue P i g ~ r i e n tof Cl~~.o?nodo?is z e b ~ n . Ibid., Vol. 26, pp. 217-
223.
Giave, C.
1916. 0117trzi~a ~ I C Z Z S ~ ) ~ I IT. ~ ~ L XII
. E ~ n h l y o l o g i c a lContlibntion a n d a
S t u d y of t h e Effect of Polli Snbstance upon Derelopment aucl
D e ~ e l o p i n e n t a l Plocesees. J o z l ~ . Xoiph., Vol. 27, pp. 413-
445, 3 pls.
PERIBROKE,
BERNUDA,
J a n n a i y , 1918.
V. J. CROZIER

A CASE O F ABNORAIAI; I S H E R I T A S C E IS
DROSOPI-IILB BIETJAKOGXSTER
A a r o s ~great linlllbers of cultures one is occasionally folllicl
which gives exceptioilal results not cxplailiable by the uinal
mode of inheritance. Althougb sllch caies do not aid in ~ u i d e r -
itallclilig genetic problems unless the ~lleclialiislli illvolved is
worliecl ont. the present case seems to be iufficiently remarlcable
to lnerit brief ~lzention. The v r i t e r has no esplallatioll to offer.
I n culture 76, xirhich was lllacle up February 9, 1917. a large
preponderance of males was obierved. the ratio being 35 lllales
to 3 felllales, and the l l l a l ~ sincluded unexpected classes. The
parents of the culture were one h011107ygous eosin ruby forlrecl
female from stock and a Illale which n a s normal wilcl-type in
all respects with the exception that the posterior cross veins of
the wings were ~llissing. 1 3 5 pedigree i\ u n l i n o ~ ~aiicl
n he lvns