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FISHERY BOARD FOR SCOTLAND.
[SCIENTIFIC INVESTIGATIONS.
LIGeLO::
No. I.
REPORT ON THE REPRODUCTIVE ORGANS OF
SPARUS CENTRODONTUS, Delaroche; SPARUS
CANTHARUS, L.; SEBASTES MARINUS (1.);
AND SEBASTES DACTYLOPTERUS (Delaroche) ;
AND ON THE RIPE EGGS AND LARVAL OF
SPARUS CENTRODONTUS (?), AND SEBASTES
MARINUS
(wir 5 Puarss).
BY
H. C. WILLIAMSON, M.A., D.Sc, F.R.S.E.
This Paper may be referred to as:
‘* Fisheries, Scotland, Sct. Invest., 1910, I. (Sept. 1911).”
TON DON:
PUBLISHED BY HIS MAJESTY’S STATIONERY OFFICE.
To be purchased, either directly or through any Bookseller, from
OLIVER & BOYD, Tweeppate Court, EpinsurGH ; or
WYMAN & SONS, Lrp., Ferrer Lane, E.C.; or
E. PONSONBY, Lrp., 116 Grarron Srreetr, Dustin.
PRINTED BY
JAMES HEDDERWICK & SONS, LTD.,
Av ‘“‘THE CrTizEN Press,” Sr. Vincent PLACE, GLAscow.
One:
Price One Shilling and Sixpence.
FISHERY BOARD FOR SCOTLAND.
ON THE REPRODUCTIVE ORGANS OF SPARUS CENT RODON-
TUS, Delaroche; SPARUS CANTHARUS, L.; SEBASTES
MARINUS (L.); AND SHBASTES DACTYLOPTERUS
(Delaroche) ; AND ON THE RIPE EGGS AND LARVA
OF SPARUS CENTRODONTUS (?), AND SEBASTES
MARINUS.
BY
H. CHAS. WILLIAMSON, M.A., D.Sc, F.R.S-E..
MARINE LABORATORY, ABERDEEN.
(Plates I.-V.)
Sparus centrodontus, Delaroche.
This species, Cuvier and Valenciennes* state, was named by
Delaroche Sparus ecentrodontus. According to the French authors,
there is doubt as to the identity of the fish named Sparus orphus, L.
Dayt gives the name Sparus orphus, Lacép, as a synonym of Sparus
centrodontus, Delaroche. Cuvier and Valenciennes proposed to change
the name to Pagellus centrodontus. But there is, in my opinion, no
necessity for removing this species and Sparus cantharus, L., from the
genus Sparus.
This species is landed at Aberdeen by trawlers during the greater
part of the year. In September, 1910, it was being captured in
especially large quantities.
It is known as the Jerusalem Haddock, Bream, Silver Haddie
(Loch Tyne).
A description of this fish is given below.
Specimens{ have been examined during each month of the year for
the purpose of studying the development of the reproductive organs.
In no instance was the ovary ripe. It seems probable that when
spawning these fishes may retire to places where they are not taken
by the trawl.
The Reproductive Organs.
This species is hermaphrodite. A pair of testes and a pair of
ovaries are present in each fish, but the testis and ovary are not
always functioning simultaneously. ‘The testis and ovary are
intimately combined on each side of the fish.
* Cuvier et Valenciennes—‘‘ Histoire Naturelle des Poissons.” T. vi. Paris,
1829.
+ Day—‘‘The Fishes of Great Britain and Ireland.” 2 vols. London,
1880-1884.
t Mr. Eunson, fish merchant, Aberdeen, has very kindly given me most of the
examples of this and the other species dealt with in this paper.
(4116) Wt. 403/46—500—9/1911,
4 Fishery Board for Scotland.
While in some fishes both the testis and the ovary may be found
developing, in others only the testis or the ovary is developing. In
the latter case the portion of the organ belonging to the other sex
may be so small that it may escape notice ; but, so far as my experience
goes, it is present. All are potential hermaphrodites, but only a
proportion are functional hermaphrodites.
The fishes have been arranged in the table in the following classes :—
(1) predominant males, (2) predominant females, and (3) sexes equal.
The predominant male or female indicates that the testis or the ovary
is large, while the organ of the opposite sex is very small. The
number examined during each month, with their range of length, 1s
given. The length of the fish is the distance from the anterior tip of
the snout to the extremity of the longer tail ramus.
The range in size of the ova found in the fishes is entered in the
last column.
A number of the fishes were spents.
The fishes entered opposite each month do not all belong to the
same year. The observations were carried on over a period of three
and a half years.
SPENTS.
PREDOMI- PREDOMI-
NATING NATING Equal ; j
MALE. FEMALE. Ty Predominat- Predominat-| *Range of Size of
MOonru. ing Male. ing Female.j the Ovarian Ova.
Diaineter in mm.
5 By 3 he uni Be || ee S|
=| b: [2 2| es 12] es |S] we
B| 88 | 5 EB) g8 78] 38 | 8) ge
Z| & Z Z| = Z| = z | =
vans 2 |e si 2¥/ | | BRI || BY 7 | 32-3 Sy :
Feb. 3 34-43 |) 35, 5 32-34 J1(?) 37 Ps =<
March 1 37 2B) + 38F Bh 2) Small. oe ae
April af ar 1} 38 *¢ er. v 55
May 9 | 34-47 | 14 33-48 | 1 | 33 is ai
June 6 | 32-36 9 4 1 36 1 38
July 9 34-43 | 32 6 39-39 3 34-39 3 39-45
Aug. 8 34 43 2 BB Apa Bae es oe ec
Sept. 36 | 34-50 7 | 31-40] 5) 37-46) 4] 39-46
Oct. 10 | 32-44 1 34 : Me Ts)
Novy. 8 | 34-46 il kt 3 "06; “172 nabs
Dec. 2) 34,35 5 | 35-37 12-92, *95, °3.
*The sizes given include all the samples from all the groups that were measured during the month.
Sample eggs from every ovary were not measured. But the ovaries were examined for any indication
of ripening ; it is not therefore likely that in any of the fishes there were eggs of greater diameter than
“45 mm.
Out of 404 fishes, 38, measuring from 31 to 40 em., had the sexes
equal, while 128 predominant males measured from 31 to 50 em., and
238 predominant females measured from 32 to 49 cm. It may be
noted that the fishes having the sexes equal did not appear among the
specially large individuals.
A number of reproductive organs were obtained from a collection
of fishes captured off the west coast of Orkney in January.
The hermaphrodite condition is indicated when a testis and
an ovary are combined in one organ, ‘They may be of equal
size, as, for example, in Fig. 21, or the testis or the ovary may be much
the larger, as, for example, in Figs. 3a and 19 respectively. In the
latter division the very small sexual organ seems to be quite normal,
and is, I consider, functional. These are the conditions where the
sexual organs are not specially large, ¢.c., in small bream (e.g., Figs.
3a, 9, 19, 38, which are of natural size). But when one sex pre-
Reproductive Organs of Sparus Centrodontus. 5
dominates in the reproductive organ of a big Centrodontus, the other
sexual organ is so much reduced as hardly to be recognised ; it seems,
indeed, to be exhausted. This condition is seen in Fig. 40 (natural
size), where, while the ovary is large, the testis is reduced to a thread.
In the previous case the fishes were, I think, preparing to spawn for
the first time. In the latter case the ovary had been ripe before.
The conclusion, then, is that the organ when it first ripens is
hermaphrodite ; at the next spawning it may be unisexual. I do not
know whether the hermaphrodite condition may persist in the second
spawning. It seems not improbable that where in the hermaphrodite
condition one sex specially predominates, that will be the sole function-
ing sex at the next spawning.
Organs in which two sexes were equally represented are given in
Figs. 17 and 21. The testis was white, ripe. It was possible to make
out the heads of the sperms, but not their tails. The ovary had an
amber tinge. In one specimen, Fig. 21, the eggs measured up to
‘Imm. in diameter, while in the other they reached a diameter of
‘15mm. A section of the latter reproductive organ is given in Fig. 18.
Where the male predominated the testes were nearly all large and
white. In one case the testis was ripe. It was pasty in consistency ;
the heads of the sperms, but not their tails, were made out. The ovary
was very small and colourless ; it was enclosed in a deep groove in the
edge of the testis. The ovary was 4mm. broad, and it was sunk flush,
almost out of sight. In Fig. da the groove has been opened out to
expose the ovary (ov.). It is quite a normal immature ovary, with
ridges filled with minute eggs, measuring about ‘05mm. in diameter.
In another fish the ovary was only partly sunk in the groove (Fig. 9).
The eggs measured from -05mm.—"07mm., in diameter. They were
colourless, non-yolked, and clear. These ova have a distinct round
nucleus.
Sometimes the ova ‘05mm. in diameter, although translucent, are
not perfectly clear, as the substance of the egg is granular; they have
an exhausted appearance.
The dorsal mesentery (dm., Fig. 38), which supports the organ to
the roof of the abdominal cavity, runs along the ovary. The testis
forms the free ventral border of the united organ. Sometimes the
side of the ovary is attached to the testis; in other cases the attach-
ment is very slight (e.y., Fig. 25).
A section across a predominating male organ, at a quarter of its
leneth from its posterior extremity, is shown in Fig. 8. The ovary is
a sac, on the inside wall of which are arranged ridges filled with ova.
The ridges run longitudinally. In some cases, ¢.g., in a fish 43°5cm.
long, the egg-ridges rose little, if any, above the ‘surface of the ovary.
The eggs were clear and nucleated, the largest measuring ‘06mm. in
diameter. Part of the inside wall of the ovary is without ova, 7.e., the
oviducal part.
The ovary and the testis do not communicate. Between the two
there is a layer of lacunar, or spongy honeycomb tissue, which extends
into the wall of the ovary (sp., Fig. 8). This lacunar tissue acts as the
vas deferens.
The testis is of an honeycomb structure. A surface view ofa portion
of the ripe testis is shown in Fig. 16. It is seen from the outside.
The spaces are occupied with sperms. The internal structure of a
whitish developing testis consists essentially of tubules (¢f, Fig. 11).
6 Fishery Board for Scotland.
This drawing represents a transverse section of a portion of a testis.
The tubules, which are arranged in a columnar form, are in some
cases branched, and they appear to connect directly with the series of
ducts occupying the core of the lobe. Each tubule is enclosed at the
surface of the testis in a loculus formed by partitions arising from the
outside wall (sk., Fig. 10). The skin of the testis when torn tends to
split longitudinally. |
Some testes were mottled. The yellow mottling is seen in the core
of the testis (mt., Fig. 24). May this mottling indicate that the fish
has spawned before ?
The egg-ridges vanish in the posterior part of the ovary, which
becomes the plain-walled oviduct (Fig. 59). This figure represents a
view of the two organs in a predominating female. The ovaries are
large, and the testes (7’) are very small. On the side of the ovary next
the wall of the abdomen there is a digitiform region of the wall of the
ovary without egg-ridges. It is the oviduct (ovd.).
The testes do not unite.
The general arrangement of the organs in the abdomen of a
predominating male is shown in Fig. 6. The external aperture of
the vas deferens is omitted.
A series of transverse sections, not drawn to scale, illustrate the
arrangement of the parts. Fig. 24 is a section of a functional male
organ near its anterior end. The ovary has only a small attachment
to the testis. A section made near the Junction of the ovaries exhibits
a similar condition (Fig. 25). Large lacune are visible in the wall of
the ovary. Egg-ridges are present, but not on the portion of the wall
next the lacune. In a section further posteriorly, but where the
ovaries are still separate (Fig. 26), the lacunar tissue in the wall of the
ovary is very extensive. Ova are present on part of the wall of the
ovary. The testis had already ended at this point. A small lobe was,
however, found in a section of the single oviduct (Fig. 27) posterior
to the union of the ovaries. The junction of the ovaries is shown in
Fig. 28. In the section of the oviduct (Fig. 27), the lacunar tissue in
its wall represents the two vasa deferentia, which it was not possible
to separate. The lacunar tissue is apparently present throughout all
the wall of the oviduct. Further posteriorly the lacunar tissue is very
much increased (Fig. 32). It is present in the whole of the wall of
the oviduct, the mesentery forming a break or division. The lacunar
tissue was broader on one side than on the other, but that may be
simply due to unequal contraction. The strands which form the net-
work of the lacunar cavities appear to be muscular. A surface view
of the inside of the last portion of the oviduct is shown in Fig. 35, .
The duct has been slit open along its dorsal (7.e., posterior) side. The
lacunar tissue can be seen in parts through the lining skin of the
oviduct. It has a different appearance in different parts.
Near the end of the oviduct the lacunar tissue (la.) forms very
large cavities (Fig. 33) on the dorsal side, and in a section made close
to the external aperture, 7.¢., next the ventral wall of the abdomen,
spaces are seen to have grown into a definite tube on the dorsal edge,
the vas deferens (VD., Fig. 34). The posterior mesentery cuts the
lacunar tissue into two.
The further relationship of the parts is shown in a longitudinal
section through the cloacal region (Fig. 31). It is shown upside down.
The lacunar tissue on the posterior side is seen to open out into a
Reproductive Organs of Sparus Centrodontus. 7
wide vas deferens, which finds exit on the top of the urogenital papilla.
The opening of the vas deferens is immediately in front of that of the
urethra (wrth.), the two apertures being separated by a thin, soft
septum. On the anterior side the outer and inner skins of the
oviduct wall join together to close the lacunar space, which com-
municates round the neck of the oviduct.
On the end of the urethra there is a bulbous gland-like swelling.
When the papilla (pap., Fig. 29) is examined from the outside, the
single crater in its tip is seen to be divided by a transverse septum.
The papilla is not large; it is larger in the male than in the
female; sometimes it is inconspicuous. It is well supphed with
blood, and is formed of very fine spongy tissue. The sides of the
crater are sometimes soft and gauzy ; this condition makes it difficult
to separate the two small apertures.
The oviduct (ovd., Figs. 29, 31) opens in front of the vas deferens,
between it and the anus. The aperture appears to be partly closed
sometimes by the cohesion of its soft lips. The urogenital papilla
and the anal and oviduct apertures are sunk in the cloacal slit. The
slit is longitudinal and it closes neatly.
When the female sex predominated the ovary was large, and pink in
colour. The testis was small, in some cases so reduced as to be hardly
visible. The ova, when “3mm in diameter, were just visible to the
naked eye. When magnified they were opaque, and were enclosed in
a swollen follicle (Fig. 20). Some eggs from another ovary
measuring ‘*3mm in diameter were not quite opaque. The nucleus
(germinal vesicle) was visible, and it was pigmented. Eggs which
are storing up yolk have orange-coloured pigment in them. The
pigment is located in the nucleus as a rim of oily opaque corpuscles
(pi., Fig. 44). The size of the germinal vesicle varies (cp. Fig. 51).
It is possible that it has the power of expanding and contracting.
‘The clear eggs measured ‘15mm. in diameter. A few specks of red
colouring matter were made out in some clear eggs of that size.
The general arrangement of the organs in the abdominal cavity is
shown in Fig. 2. The ovary was large and orange-coloured. ‘The
testis (7') was rendered visible by some brown mottling along its
extent. The colouring matter is in little traces as well as in com-
paratively big patches.
A fine yellow mottling is to be seen scattered through some of the
ovaries. The ovarian ridges on the oviduct region of the ovary are
covered with a delicate cuticle, which can be easily dissected off in a
piece.
The mesentery (dm., Fig. 40) attaches the ovary to the swim-bladder.
The principal blood-vessel is situated on the side facing the other ovary.
The ovaries unite posteriorly, and end shortly thereafter. The
longitudinal egg-ridges, seen in section, tend to radiate from the large
blood-vessels. On the ridges are developed blunt villiform processes
full of ova. A developing ovary, which had been a spent, is
shown in Fig. 40. The testis, which is a very narrow fold, is in some
parts flush with the skin of the ovary. It could be traced right up to
the anterior end of the organ. The oviduct region (ovd.) is well
marked. In some cases, however, the testis began some distance short
of the anterior end of the ovary (cp. Fig. 22).
The testis is sometimes hardly visible; that appears to be the case
especially in ovaries that have been ripe before. The testis in Figs.
8 Fishery Board for Scotland.
19 and 38, although small, is functional and readily seen, while in
Pig. 40 the testis is so much reduced as to be probably non- functional, ;
and might be easily passed over. In the last ovary old egg- -capsules
were present, the remains of a previous spawning. The two classes of
reproductive organs differ also in the amount of empty space in the
early developing ovary (cp. Figs. 42 and 45).
Three sections were made across the organ shown in Fig. 40, viz.,
at the points A, B, and C. ‘The point “A is near the anterior ex-
tremity: the section is given in Fig. 45. The testis is visible as a
little projecting fold: the oviduct is absent. No part of the wall of
the ovary is free from eggs, but the ridges are narrow on the dorsal
and ventral sides. In a section at B, viz., across the beginning of the
oviduct, Fig. 46, the oviducal part is very narrow internally ; there is
little, if any, clear space. The testis is flush with the outer ‘skin. The
ovary has a big lumen. In the posterior part of the ovary at CO, the
oviduct is broader ; the testis is a little fold (Fig. 47). Near the union
of the ovaries, ae at D, the outside breadth of the oviduct is not
equalled by plain surface on the inside. The plain surface of the
oviduct inside an ovary is shown in Fig. 23. It exhibits a quantity
of minute white granules, which are thickly arranged about the
middle and gradually get fewer towards the egg-ridges at the sides.
Minute muscle-fibres are visible in the wall.
In the small ovary, Fig. 38, which is probably ripening for the first
time, the oviduct can be traced further anteriorly than in Fig. 40 (cp.
the cross sections, Figs. 42 and 43).
A section across the union of the ovaries of another fish shows the
posterior end of the septum (Fig. 49). In this case the testes were
dorso-lateral in position, while in another example they were ventro-
laterally placed. At a short distance posterior to the above point,
Fig. 50, the wall of the oviduct was spongy, thick, extensile, all round
except at the ventral region. The testis was pr esent on one side only.
The oviduct is wider than in the male. On the last part of the
oviduct I observed two lacunar areas, one bigger than the other, with
intervening plain areas, which exhibited straight longitudinal fibres.
The external opening of the oviduct is large; the lips are some-
times broad and leaf-like.
The urogenital papilla in the female is small. The opening on its
tip is crater-like, and on first examination it appears to be a single
aperture. But if a section be made across the papilla the two ducts
are found. The urethra has a thick wall, and its lumen diminishes
somewhat near the aperture. The vas deferens is a slit which is
closed (Fig. 41). The two ducts meet just at the aperture. The
papilla is formed of spongy material.
It is an interesting problem whether a fish which becomes unisexual
will remain always of that sex. It seems probable that it will do so,
for in several developing predominant females I have found the old
capsules of ripe eggs which had been retained in the spent ovary.
(Vide also p. 10.)
Notes on the Reproductwe Organs.
The stage of development of the ovary in the fishes examined in
each month is indicated by the size of the eggs entered in the last
column of the table on page 4.
Reproductive Organs of Sparus Centrodontus. 9
In January, February. March and May some of the testes were ripe.
In June, July, and August, while there were small ovaries and
testes, some spent ovaries were found.
In September the fishes presented large and small testes, developing
ovaries, and spent ovaries and testes.
In October the testes and ovaries were small and developing; one
ovary was spent.
In November fairly large testes and ovary were found. The fishes
had much abdominal fat.
In December a small testis was growing white; an ovary was
developing. A considerable quantity of abdominal fat was present.
The Ripe Kggs.
No ripe ovary was obtained, but in several fishes the old capsules
of ripe eggs were still present. The eggs were crushed and dried.
Only a little white amorphous matter remained inside. An opaque
circular mass, sometimes stained yellow, could be detected in the
amorphous material. The yellowish mass measured about ‘27 and ‘3
mm. in diameter. In two cases when the egg was dissected a little
white coherent irregular body, measuring about 2 mm. in diameter,
was found. This, I think, may be the remains of an oil globule. In
September a quantity of unspawned crushed ova remained in an ovary.
A bright golden oil globule was present in the ege. In one it
measured *25 mm. in diameter.
The crushed capsules were oval and the eggs had evidently been
oval. Several of these capsules were measured; the sizes were as
follows: —1°35 x°8, 1:°2x°75, 1:2x-75, 1:2x°75, 1:1x-9, 10x89.
1:0 x ‘7 mm.
Undeseribed Pelagic Fish Egg.
An egg which appeared in the surface tow-net collections made in
1903 by the Fishery Board, in connection with the International
Investigation of the North Sea, was remarkable in that it was oval in
shape and had an oil globule (Fig. 13).
The eggs were measured after preservation in formaline solution.
The oil globule varied from ‘17 to ‘12 mm. in diameter. The following
are the sizes of the eggs in which the oil globule measured ‘17, alo
‘12mm. respectively :—Oil globule -17 mm.—eggs, 1-4 x PP2E 37 Xx
11, 1°32 1:17, 1:31:05 mm. Oil globule 15 mm.—eggs, 1°37 x 1+1,
1°35 x 1-2, 1°35 x 1-07, 1-35 x 1:05, 1:32 x 1-1, 1:32 x 1-07, 1:32 x 1-05,
1°3 x 1:07, 1-27 x 1:1, 1:25 x 1:1. 1:25 x 1:07mm. Oil globule -12mm.—
egg, 1°35 x 1:1 mm.
I consider that this egg is that of Sparus centrodontus. It was cap-
tured in May at points about 40 miles east and 15 to 20 miles west
of Shetland respectively. The embryos were all early, except one
which extended for about two-thirds round the yolk. One egg was
obtained in June, about 20 miles west of Shetland. The embryo was
in the disc stage. In August a large number, 84 in all, was obtained
off Kinnaird Head, Moray Firth. The exact position was—Troup
Head, S.W. by W.; Kinnaird Head, S.E.3S. The eggs were all
recently spawned, the embryo being in the disc stage. A very deep
hole (100 fms.) occurs in this vicinity, about six miles off-shore,
10 Fishery Board for Seotland.
During the same month one egg was got between Orkney and Shet-
land, viz., ten miles E. of Fair Isle. The embryo was about three-
quarters developed. Thirty-eight eges were captured 40 miles E. of
Shetland in this month. The embryos were about ready to hatch
(Fig. 5). The embryo dissected out of one of the eggs is shown
enlarged in Fig. 12.
Almormal Reproductive Organs.
A fish measuring 47 cm. in length had a normal reproductive organ
on the left side (/.) and an abnormal organ on the right side (Fig. 1).
The latter contained several dark-coloured concretions(c.), while a small
concretion (¢’) was attached to the testis of the left organ. The
concretions are collections of unspawned eggs. The concretion ¢”' is
hard and dark-coloured. It is situated in the ovary. It cuts like a
hard cheese, and shows in section a conglomerate of eggs. The eggs
in the lighter-coloured balls, ¢. and ¢’, appear to be formed of the eggs
of a later spawning than ec”.
The quashed eggs measured 1:5 mm. in greatest length. The cap-
sules were flattened and the yolk had disappeared from many alto-
gether, abhoueh still present in some. ‘The old eggs had a single oil
globule :22 mm. in diameter.
The nate comb structure of the testis was well seen in the right
organ. Clean oval holes were observed in the septa of the honeycomb
tissue.
The lett ovary was developing, while the other remained in a state
of quiescence. The presence of the ball of unspawned eggs had eyvi-
dently inhibited the development of a new crop of ova. Will the for-
mation of the new ova be prevented if the ovary is in a state of com-
pression ?
In another fish the anterior portion of the ovary was detached from
the remainder (Fig. 19). No testis was observed attached to the isolated
portion.
A Mish with a Second Anus.
A fish which measured 42 cm. in length had a second anus. ‘The
gut was attached to the wall of the abdomen, which was perforated by
a pore that opened into the gut (Fig. 3).
The usual condition of the gut is as follows :—After leaving the
stomach the gut (d) passes back to the end of the abdominal cavity ;
it then turns round and comes forward to the anterior end of the
cavity. It then returns posteriorly and proceeds directly to the anus.
The three portions of the gut are looped together by mesentery. The
middle portion of the gut is the part which was connected to the second
anus (av). An enlarged drawing, Fig. 7, shows the gut and the part
of the abdominal wall (1). A fat-fold is attached to the mesentery.
Two encysted parasites (c) were located in the mesentery. The con-
tinuation of the gut through the body wall is shown in Fig. 15. The
antero-posterior direction is indicated by the letters a—-p. The
secondary anus was 2mm. long. ‘Two little processes were present on
one side (Fig. 14).
It seemed as if very little matter could have found exit by this
aperture.
Reproductive Organs of Sparus Centrodontus. 11
DESCRIPTION OF Sparus centrodontus, Delaroche.
This fish is coloured a bright red or pink. Over the abdomen there
is a golden red sheen. Small specimens show little red on the body ;
they are more silvery. The snout, cheeks, and sides are silvery with a
golden tinge. A prominent black patch is present on the shoulder,
on the beginning of the lateral line. The lateral line is prominent.
The dorsum has a metallic appearance. The top of the head is
pinkish. The inside of the mouth, and the inside surface of the gilJ-
cover, are pink and orange-red respectively.
The scales are ctenoid (Fig. 60); they are large and hard. The general
arrangement of the scales is indicated in the drawing Fig. 4; the
scales themselves are represented diagrammatically. ‘The scales are
arranged in parallel rows. This gives rise to the longitudinal rows
(or lineation) which are made out on the side of the fish. The large
semi-circular scale situated immediately over the opercular cleft is note-
worthy. Alongside the base of the ventral fin one or two scales are
elongated into a sharp triangular form, which resembles a short spine.
The scales are, however, thin and flexible. On the ventral surface
the scales between the ventral fins are extended posteriorly in a
triangular projection.
All the fins are red. The exact number of fin-rays is not entered
in the drawing. The number of fin-rays in the dorsal and anal fins
is shown in the following table :—
[
Dorsal. Anal.
Length
of Fish. a4 Soft A AS Soft
PCE Te Raye. SPMes hRaws
em. E
36 (25 | Fin-rays). 3 3
37 Aiea 42 z 18
Boe Wil 18 3 15
42 toes | 12 3 | D
4,2 12 | 15 3 | 3 |
45 12 13 3 Pein |
46 12 13 3 Ae
The dorsal fin is composed of spines and soft rays. Of the former
there are usually 12, and of the latter 15. The first of the soft rays
ends in a single tip as a rule; in one fish the tip was split. The
remaining soft rays are split. The dorsal fin stands in a groove in
the dorsal edge of the body. The front halt of the fin, consisting of
the spinous rays, can be laid down into the groove. The last two
rays are close together at their bases. The first two spines and the
last two rays arise from a single interspinous line in each case. The
lip of the hind portion of the groove is formed by large thin scales
which rise up from the dorsal edge on each side of the fin. The
scales are continued upon the bases of the last two rays as a process
similar to that shown for Cantharus in Fig. 113.
12 Fishery Board for Scotland.
The anal fin, like the dorsal, stands in a groove, the lip of which is
formed posteriorly of scales which are continued in a process upon
the proximal part of the last tworays. The first three rays are spines.
The last two rays are close together at their bases. The first two
spines and the last two rays arise from a single interspinous bone in
each case. ‘l'he angle between the last ray of the dorsal and anal fins
respectively and the edge of the body is not filled up with a
membrane.
The pectoral fin has 16 rays.
The ventral fin has 6 rays; the first of these is a spine. Only a
very small portion of the angle between the last ray and the ventral
surface of the body is filled up with a membrane.
The anus and the apertures of the genital organs and urethra are
in a slit-like depression on the ventral edge—the cloacal slit. It is
indicated at a.
The teeth in a centrodontus 40 cm. in length were small. They were
of two distinct kinds. On the premaxille there were at the
symphysis about four rows of small curved sharp teeth. Posteriorly
there were three rows; of these three the outer row were tusk-like,
while the inside row consisted of rounded-topped molar-hke teeth.
In the lower jaw (Fig. 37) two rows of teeth were made out in the
dentary at the symphysis; the outer row of curved teeth, the mner
row of teeth with blunted tips. All the teeth were small, but the
outer row contained the largest. Posteriorly the inner row showed
the rounded molar-like teeth. .
The upper lip is papillated ; the papillee dip in between the teeth
of the outer row. The papillation occurs in the lower jaw only to a
slight extent.
Inside both upper and lower jaws there is a pouch formed by a
horizontal membrane joining the premaxille and dentaries respectively.
A longitudinal strip of red muscle is present on the side just
beneath the skin.
The peritoneum is black.
The urinary bladder is in a little chamber cut off from the end of the
abdominal cavity by a septum of peritoneum crossing the cavity
between the oviduct and the urinary bladder. The chamber is lined
with peritoneum. The urinary bladder is wide; it is sometimes
covered with a layer of fat. A mass of fat is sometimes found in the
posterior end of the abdominal cavity. The ureter (ur, Fig. 6)
comes down medianly in front of the first hamal spine, and crosses
over the left side of the swim-bladder to join the urinary bladder.
The swim-bladder is large. A thick yellow matter was found in
the hind end of the swim-bladder of a fish 46 em. long in November.
Posteriorly the swim-bladder is firmly attached to the first heemal
spine. The first heemal spine ends the abdominal cavity and also the
swim-bladder. The first and second hemal spines touch one another.
The interspinous bones are long (int., Fig. 2).
The number of vertebrae in one specimen was 23. The first heemal
arch was on the tenth vertebra.
Sparus cantharus, L.—The Black Bream.
This species is landed by trawlers at Aberdeen from time to time.
One example examined in November in the fresh condition measured
Reproductive Organs of Sparus Centrodontus. 13
40:5 cm. in length. It was a silvery fish, with a slight bluish tinge
on the dorsum. The cheeks were silvery. The top of the head was
of a purplish colour. The inside of the mouth and the inside surface
of the gill-cover were white.
The lateral line is prominent (Fig. 30). The scales, which are
ctenoid, are large, and are arranged in parallel rows. A series of
longitudinal rows is prominent. A golden sheen is observed in the
rows sometimes. The large semi-circular scale situated immediately
over the opercular cleft is noteworthy.
The mouth is small, with prominent lips; it resembles the mouth of
Labrus. The teeth are comparatively large. They cover a broad
area in the jaws (Fig. 36). There are at least six rows of teeth in the
front of the lower jaw. The outside row of teeth are the largest.
They are sharp-pointed. The teeth gradually decrease in size towards
the inside of the jaw. No round-topped molar tooth was made out.
There is in both jaws a pouch formed by a horizontal membrane
joined to the premaxille: and dentaries respectively.
A strip of red muscle is present along the middle of the side.
The peritoneum is white.
The urinary bladder is large. Its attachment in the hind end of
the abdominal cavity is median, but it extends forward on the right
side of the body in the form of a broad lobe. It reaches to the
rectum. It is very vascular.
No urogenital papilla was observed. The apertures of the urethra
and vas deferens were sunk in three cases examined.
All the fins are dusky in colour. The fins are composed of spines
and soft rays.
The dorsal fin is dark, but the web between the rays was blotched
with pink. There were 11 spines and 13 soft rays in this fin in three
cases, 10 spines and 14 soft rays in one case, and 11 spines and 12
soft rays in another case. The first of the soft rays had a very fine
stiff but flexible point. The last two fin-rays arise close together.
This fin stands in a groove. The first half of the fin can be laid
down and almost hidden in the groove. The hind part of the groove
has thin, soft edges formed of scales, and these run up on to the last
rays, in a free process (Fig. 113). A similar arrangement is present
in connection with the anal fin, but here there are two processes on
each side, one a little anterior to the other. The anal fin had 3 spines
and 11 soft rays in five cases. The first two spines arise from the
first interspinous bone. The last two rays arise close together from a
single interspinous bone. This fin had its base, both fin-rays and
web, whitish ; its distal half is blackish. In one fish the fin was
bluish. The anal fin, all but its last three rays, stands in a groove.
When the dorsal and anal fins are distended the spinous and soft-
rayed portions of the fins are not marked off from one another. The
fin has the appearance of a single fin, not of two fins joined
together.
The pectoral fin is almost colourless. It had 16 rays, and reached
to the level of the anus in one fish.
The ventral fin is dusky ; the rays were bluish in one. It had six
rays. The first ray is a spine. Alongside the base of this fin an
elongated but thin scale is present. At the ventral surface between
the ventral fins the scales project in a triangular process. In a fish
42 cm. long the dorsal ramus of the tail was the longer.
14 Fishery Board for Scotland.
The number of vertebree in three cases was 24. ‘The first hamal
spine was on the eleventh vertebra.
The otolith is convex on one side and concave on the other. Views
of both sides and also an edge view of the otolith are given in
Figs. 52, 53, and 54.
Comparison between Centrodontus and Cantharus.
The two species may be confounded. They resemble one another
in the general shape of the body. Ifthe upper jaws of Cantharus be
retracted and the mouth shut, a specimen of this species might be
taken for a pale-coloured example of Sparus centrodontus. Some-
times the smaller Centrodontus (e.g., 20 cm. long) show little red
colour, and they may be mistaken for Cantharus, i.e., when the colour
is taken as a guide.
The longitudinal rows are rather more marked in Cantharus. The
eyes of that species are smaller than those of Centrodontus.
Cantharus has a smaller mouth, and more and larger teeth, than
Centrodontus. They both have a prominent dorsal ridge in front of
the dorsal fin: that of Cantharus is the more acute. They differ in
the shape of the head, seen in profile. Centrodontus has a more
rounded and fuller snout, but the difference is not always easily
ganged.
Cantharus has less body thickness, from side to side, than Centro-
dontus.
The two species differ in the colour of the inside of the mouth, gill-
cover, and peritoneum ; vide pp. 11 and 13.
The Reproductive Organ of Sparus cantharus.
The reproductive organ resembles that of centrodontus ; it is herma-
phrodite. Ina fish 40°5 cm. long, a predominant female, the ovary was
orange-coloured. A little testis was present along the lower border
of the ovary. The ovarian folds filled up the ovary (Fig. 48). These
folds are composed of digitiform processes filled with eggs. The
organ does not appear to have been ripe before. The yolked eggs
measured *35 mm, in diameter. Most of the eggs were small and
clear. There was a slight pink sheen in the yolked eggs when they
were examined with the microscope. The egg filled the follicle
almost completely. There were little masses of opaque yellow
pigment throughout the ovary.
_ A fish 42 cm. long wasa predominant male. The testis was small
and mottled. It had, I think, been spent. The ovary was a wide
sac. Some small eggs were seen in the wall of the ovary. The
lacunar tissue in the wall of the ovary was very open. The external
aperture of the oviduct was closed. The urethra and vas deferens
opened by the same external pore. In a second predominant male,
39 cm. long, the ovary was a wide sac, and a quantity of yellow
glandular-like bodies projected from the internal surface. In the
anterior region they were located all round the ovary, but in the
remainder of the organ they were restricted in distribution; in
the posterior part they were merely dorsal in position. They were
arranged in a manner generally resembling the distribution of the
Reproductive Organs of Sparus Centrodontus. 15
ege-ridges, Small eggs are associated with these yellow bodies.
They are evidently the yellow giands that are seen in the developing
ovary.
The January fish 42cm. long had yolked eggs ‘37 mm. in diameter.
Sebastes marinus (1).
According to Cuvier and Valenciennes,* Linneeus included this fish,
and Perea seriba, L., under the name Perca marina, L. Miiller named
it Perea norvegica. The two French authors said that ‘its form is
nearly that of the Perch or of the large Serranus;” and further, ‘ it
is not a Perch, but a Sebastes similar to the form described by
Delaroche under the name NScorpena dactyloptera.” Cuvier and
Valenciennes gave it the name Sebastes norvegicus. Smitt, in
‘ Scandinavian Fishes,” uses the name Sebastes marinus (L.).
This fish is undoubtedly a Perca, It resembles Perea fluviatilis
very much in general shape, colouration, and external characters.
The chief difference lies in the dorsal fin, which is here a single long
fin having two distinct parts. In Perea fluviatilis these two parts are
separate fins. The exclusion of Sehastes marinus from the genus
Perca does not appear to me to have been justified. The question is,
however, complicated by the other members of the genus Sebastes,
and especially by Sebastes dactylopterus, which is described in another
part of this paper.
This fish, which is known as the Norway Haddock, or Runkie, is
landed regularly by trawlers at Aberdeen. Large quantities are
brought from Iceland, but some are also got in the North Sea.
Among the adult specimens examined there was a great difference
in size, so much so that there appeared the possibility that two species
were included under the same name. )
FIGs.
Fic.
biel
S35
23.
24.
25.
26.
27.
28.
29.
30.
Sl.
32.
30.
51.
52.
53.
54.
56.
57
59.
60.
61.
Portion of oviduct-wall of ovary of a fish 46em. long. | November.
Enlarged.
Section of testis of fish 43cm. long. November. Enlarged.
99 99 99 99 9 >
99 I) bh) ” ” 9
Section across the oviduct of fish 43cm. long. November. Enlarged.
Section across the union of the oviducts.
The apertures in the cloacal slit of a predominating male 46cm. long.
Enlarged.
Sparus cantharus, Linn. 40°5 em. long. :
Longitudinal section through the cloacal slit of a predominating male.
Enlarged.
Section across the oviduct a little posterior to the position of Fig. 27.
Enlarged.
Section of oviduct of predominating male 36cm. long. November.
Enlarged.
Section of the same oviduct as Fig. 33, but close to the abdominal wall.
Enlarged.
Oviduct of the same fish as Fig. 24, opened at its external aperture.
View of internal surface. Enlarged.
Drawing showing dentition in lower jaw of Sparus cantharus. Approxi-
mately natural size.
Lower jaw, to show dentition. Approximately natural size.
Reproductive organ, about natural size.
Reproductive organ. January. Dorsal or posterior surface.
Spent ovary developing, of fish 44cm. long. November.
Horizontal section of urogenital papilla of a predominating female,
42‘5cm. November.
Section at A of reproductive organ shown in Fig. 38. Enlarged.
>
Lasoo ae? Ree So a ”
Diagram of developing egg: p7. indicates pigment in the nucleus. En-
larged.
Section at A of the reproductive organ shown in Fig. 40. Enlarged.
99 B 99 9 O') be)
C bie)
99 3 9 be)
Section of the reproductive organ of Sparus cantharus, about the middle
of its length. Enlarged.
Section of the reproductive organ of a fish 46cm. long across the
junction of the ovaries. Enlarged.
Section of the same reproductive organ as Fig. 49, but posterior to the
junction of the ovaries. Enlarged.
Diagrams of early developing eggs, showing different sizes of nuclei.
Views of concave surface of the otolith of Sparus cantharus. Nat. size.
Edge view of the otolith of Sparus cantharus.
View of convex surface of the otolith of Sparus cantharus.
Sebastes marinus, female, 37 cm. long. Reduced.
and 58. Sebastes marinus. Solidified albuminous matter from surface
of the wall of the ovary. Enlarged.
Sebastes marinus, section of pregnant ovary.
Scale of tish 38cm. long. About natural size.
Sebastes marinus, 40 em. long. Scale about natural size.
PLATE Il.
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(1312£) 403/46, 500, 4/11, BANKS & CO.LTD.
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Sebastes marinus, male, abdomen. Nat. size.
female “8 re .
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and 65. Sebastes marinus, developing eggs. April.
Sebastes marinus, female, urinary fringe. Enlarged.
93
9
9
side view of head of embryo. May. The tail of the
embryo nearly reached the head. Enlarged.
larva, just hatched, abnormal. Enlarged.
head of larva, recently hatched. Enlarged.
developing embryo. The preserved egg measured
125mm. April. Enlarged.
female, 34°5cm. Developing embryo. May. Enlarged.
section of ovary. September.
pregnant ovary. Nat. size.
developing yolked egg, °37 x 3mm. September. En-
larged. ;
female, spent. Urinary papilla. Enlarged.
post-larva, 6°25mm. long, from ovary. June. En-
larged.
larva, length 49cm. May. Enlarged.
post-larva, 7 mm. long, from ovary. June.
small egg, about ‘4mm. in diameter, becoming yolked.
January.
portion of ripe ovary. April. Enlarged.
*
PLATE til.
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Sebastes marinus, portion of a large ovary. February-March. Enlarged.
s5 ss
spent ovary. September. Ovarian wall slit open.
spent ovaries and external apertures.
portion of ovary. April. Showing very large follicle.
Enlarged.
spent ovary, developing. April. Enlarged.
- a September. Enlarged.
male, urogenital papilla. February.
section of urogenital papilla of male. February.
Enlarged.
young egg in wall of vessel in ovary. October. En-
larged.
developing egg in ovary. October. Enlarged.
pe) eggs 9 2
portion of spent ovary. September. Enlarged.
longitudinal section through the anal region of
female. Enlarged.
dorsal fin, last interspinous bone and last two fn
Enlarged. Semi-diagrammatic.
anal fin, last interspinous bone, and last two rays.
vertebra beari ing the first hzemal spine.
dorsal fin, end of, showing membrane joining the last
fin-ray to the dorsal edge of body. .
Sebastes dactylopterus, 41cm. long. Scale. Nat. size.
9 be)
view of adnate ventral fin.
theoretical section of egg-column of ovary.
portion of ovary. January. Enlarged.
view of top of head, showing the interorbital and
adjacent regions.
egg beginning to store yolk. Semi-diagrammatie.
Enlarged.
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106.
107.
108.
LO9.
110
AGW E
112.
113.
114.
115.
PLATE V.
Sebastes dactylopterus, portion of ovary, cleared with solution of NaOH.
March. Enlarged.
= me x1. The scales are not shown in correct size.
5 a transverse section of ovary. January. Enlarged.
. 5 ovary. January. Nat. size.
SS 35 transverse section of ovary, showing the granular
matter attached. February. Enlarged.
A and B. Sebastes dactylopterus, ventral views of the anal regions of
two fishes.
Sebustes dactylopterus, ovary. February. Nat. size.
= testis. January. Nat. size.
Sparus cantharus, end of dorsal fin, to show process formed by scales.
Sebastes dactylopter us, head seen from above.
Sebastes marinus, head seen from above.
PLATE V.
H.C.w.
(1312€), 403/46. 500. 2/11 BANKS & CO.LTO.
a 3 | ‘4
FISHERY BOARD FOR SCOTLAND.
SCIENTIFIC INVESTIGATIONS,
iS TO;
No. IL.
THE RETARDATION OF THE DEVELOPMENT OF
THE OVA OF THE HERRING
(witH 1 Puate).
BY
H. C. WILLIAMSON, M.A., D.Sc., F.R.S.E.
This Paper may be referred to as:
“Fisheries, Scotland, Sci. Invest., 1910, Ul. (Sept. 1911).”
LONDON:
PUBLISHED BY HIS MAJESTY’S STATIONERY OFFICE.
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FISHERY BOARD FOR SCOTLAND.
REPORT ON THE RETARDATION OF THE DEVELOPMENT
OF THE OVA OF THE HERRING.
BY
H. CHAS. WILLIAMSON, M.A., D.Sc., F.R.S.E..
Marine Laporatory, ABERDEEN,
(Plate I.)
CONTENTS.
PAGE
The Methods Employed, : : : : : : 3
The Rate of Movement of the Apparatus, : : 4
Table of Temperatures, etc., ; : : ; : 5
Results.—The Uncooled Spawn on Gravel, : ; : : 6
The Spawn Incubated in a Galvanized Tin, 6
Cooled Spawn, . 7
Gravel and Glass, i : 2 : : . 8
Exposure to Light: Growth of Diatoms, d : : 8
The Current of Water, ; : : , : 9
The Filter- Barrel, : : : : : : 9
Experiments with Spawn from Live and Dead Herrings :—
On Galvanized Wire Gauze and Glass Plates, . 2 9
Milt and Spawn, . J : : ; : F 10
The Crystals in the Eggs, : : 3 : ; ; 10
Recommendations, : : : : ; : 11
The experiments on the retardation of the herring, which have been
carried on at the request of the Government of New Zealand, were
continued during the spring of 1911.
The eggs were fertilized on board a fishing boat on the evening of
27th February and the morning of the 29th February. The spawn
was obtained from live herrings caught in a drift net. It was
attached to glass plates and also to coarse gravel, as it was desired to
see how the two methods compared in respect to the well-being of
the ova. The method adopted in the case of the glass plates was
similar to that described in my previous paper.* For the second
method a layer of gravel was put on the bottom of a barrel partly
filled with sea-water. Some milt was pressed out into the water, and
then some spawn was pressed out and allowed to fall on to the gravel.
The gravel was left undisturbed for some hours. Twelve hours or
more later, the barrel, on arrival at Aberdeen, was tipped up and the
gravel turned out into a tub. A considerable quantity of the ova
* ** Experiments to show the Influence of Cold in retarding the Development
of the Eggs of the Herring (Clupea harengus, L.), Plaice (Pleuronectes
platessa, L.), and Haddock (Gadus eglefinus, L.).” Twenty-seventh Annual
Report of the Fishery Board for Scotland, Part II1., 1910, p- 100,
(4189.) Wt. 10769/231—500—9/1911,
4 Fishery Board for Scotland,
was not attached to the gravel. Some eggs had been killed. A fair
amount of ova was still adhering to the stones. They hatched out
well, as did some of the eggs which were found loose among the
gravel. The spawn was divided up. Part of both lots was cooled,
while the remainder was kept in uncooled water. The glass plates,
both cooled and uncooled, were made to revolve. The spawn on
gravel, both cooled and uncooled, was put into trays, which were
moved vertically. The revolution of the plates, and the raising and
depressing of the trays. was intermittent, not continuous. The trays
were made of wood, and they had bottoms sparred with glass tubes,
and in one case covered with old galvanized wire netting.
The uncooled glass plates were put into the apparatus used in the
1910 experiment. * It consisted of a galvanized iron box, which
revolved inside a galvanized iron tank.
The apparatus used for the cooled spawn was of wood, most of
which was tarred. The box containing the plates was 21 inches
square and 18 inches deep (A., fig. oy, A wooden frame, which
carried a plate on each of its four external faces, revolved on iron
axles. Each plate was in succession brought to meet the inflowing
current of water.
In the case of the uncooled gravel spawn the trays were raised a
little and then depressed. The cooled gravel spawn was arranged in
trays that travelled round on a revolving frame (B., fig. 2). The
trays were 9 inches by 7 inches by 2 inches deep.
The water was filtered through a barrel filled with sand, both for
cooled and uncooled ova.
For cooling the water a series of galvanized iron pipes (#-inch wide
diameter), covered with ice, was employed.
Sea-water had been running through the pipes and the apparatus
for three months before the ova were introduced.
The spawn was put into running water by about 10 p.m. on the
28th February.
The temperature of the uncooled and cooled water, and the
quantity of water supplied to several of the lots of eggs, are shown in
the table, p. 5. The temperature in °C. was read almost hourly
during the day and night. The maximum and minimum thermo-
meters were read once, sometimes twice, a day. The temperatures
given in the maximum and minimum columns refer to parts of two
days. The quantity of water is indicated by the time required to fill
a one-pint measure (‘6 litre).
An estimate was made of the number of eggs on two plates, which
were not too crowded with ova. They contained approximately 2600
and 2800 eggs respectively.
RATE OF MOVEMENT OF THE APPARATUS.
The cooled glass plates revolved once in from 1 to 2 minutes. The
cooled gravel in trays made a revolution in from 5 to 105 minutes.
The plates in the tin revolved once in from 14 to 2 minutes. The
uncooled gravel spawn in trays was raised and depressed once in 5$
to 74 minutes.
* ‘Experiment in retarding the Development of the Eggs of the Herring.”
Twenty-eighth Annual Report “of the Fishery Board for Scotland, Part IIT., 1911.
Lhe Retardation of the Development of the Ova of the Herring.
IncUBATION OF HERRING OvA —TEMPERATURE OF THE WATER, ETC.
Number of Days.
OMADIFWNHr
|
TEMPERATURE OF THE WATER.
capacity of 1 pint (=‘6 litre),
* The quantity of the flow of water is indicated by the tiie required to fill a vessel of a
(2) FLow oF WATER.*
i Glass @) Ur |
Gravel. Pinte Tin. cooled.
Range in | Max. and | Max. and Max. and | Max. and Nos. | Tin epee
nce Min. °F. Min, “Ey | Min. °F. | Min. ee: 1 and 2. Gravel.
- ee Secs. | Sees,
3—2°6) 42, 40 43,40 | 42°5, 40 40 ae se
d4—2°6) 42, 37 42,37 | 42°5, 41 ies - 4] | 22 95
3°2—2 38, 56 40,36 | 44:5, 43 | 44 5, 43 fe | ate
4—1°6} 38, 36 40,36 | 42°5 43'5 Be 22 a)
2°5—1°6| 38, 35 40,35 | 44°5, 39 | 44-5, 39 | ea
2°6—1°6, 36, 35 DD, ap) Pele: 4h 4g: 5, rel ere jt
2°2—1:4| 36, 35 37,36 | 41°5, 40 41°5, 40 9—2] : we
2°1—1°4)35+,35] 36,36 | 39°5 39°d 9—1 | 21,30) 14,18
1:9—1°4) 35+, 35] 56,35 | 42:5, 40 | 42°5, 40 18 22 Ly pS al
1°9—1°6| 37, 35 36, 35 | £2°5,°39 | 42°5, 39 | 16, 21 Dia) |) alts?
1°9--1:4) 35, 35 35, 35 | 42°5, 39. | 42°5, 39} 16, 22 27 20
1-9—1°4| 35, 34°5| 36,35 | 41°5, 39 | 41°5, 39 | 14—26|] 22 x
‘S+—1:4| 35,34 (35:5, 34:5] 40-5, 39 | 40°5, 39 |} 13—18| 238 38
2°2 —1-°3) 36, 34°5 | 36, 34°5 10—26| 30 5
2—1°'4| 36, 34:5] 36, 35 Be ae 14—23| 25 ae
2°4—1°6| 36,35 | 36, 35°5 | 40°5, 38 | 40°5, 38|}13—18| 44 | 46
22—1-7} 36,35 | 36, 35°5 ae ie 11—27 /18—46| 10
2— 1:6} 35°5, 35 | 36, 35°5 | 41°5, 40 | 41°5, 40 | 13,17 19 P10. 29
2—1°4/35°2, 34°5) 35°5, 35 : 10—15 5 | re
1°7 — 1°4/35-2, 34°5) 35°5, 35 iB 11—16); 19 31
19—1°3| 35,35 | 35:5, 35 | 40°5, 37 | 41-5, 39 18
2—1°6| 35°2, 35 | 35°5, 35 | 36, 35 ! 13—18 ~ ae
1°8—1°6/35°2, 34°5| 35°5, 35 | 37 41°5, 40 | 14—18] 100 32
2—1°4/35°2, 34°5) 35-5, 35 | 37,35 | 39-5, 39 17 35 30
2—1°4/35°2, 34°5| 35°2, 35 | 37, 35°2 | 42 eee) 34 32
1°8—1°4/35°5, 34°5) 36, 35 ihe ae ve 16 25
1:9—1-4/35°5, 34:5] 36, 35 dt,9) (42°3, 40) 16 40 j
2—1-4| 35, 34:5 | 35°5, 35 | 37,36 | 42:5, 41) 18, 20 Se |
1‘9—1-°6} 35,35 | 36, 35°5| 37,36 | 42°5,41/ 20 65 |
1:9—1°6] 35, 34:5 | 36, 35:5 | 37, 36 ; /15—231] 55 |
2 —1-6|35-2, 34-5] 36, 35-5 | 37-5, 36 PE eo 75 |
2—1:4|35°2, 34°5) 36, 35°5 | 37:5, 36 | 43:5, 41 |16—28) 65
2—1°4/35-2, 34°5) 35°5, 35 | 37,36 | 42-5, 41 |17—26
1°9—1:2| 35,34 (35:5, 34:5) 38,35 | 42°5, 40] 15, 28 $i
1°8—1-4| 30, 34:2 |(35°5, 34-5) 37,35 | 41°5,39/13—17| 63
1°8—1°2| 35, 34:2 |35°5, 34:5) 37,35 | 40°5,39|)14—17| 63
1‘8—1:2| 35,34 |35°5, 34:5) 36,35 | 39:5, 38|}14—30| 65
1°8—1°6} 35,35 | 36, 35:5 | 37°5, 35 | 42°5, 39 | 15—30
1:9 — 1-3)35:2, 34:2} 36, 35 37, 37 42°5, 42 | 15—25
2—1°4| 35, 35 36, 35 37, 37 ine 20
1°8 —1:2/35-2, 34°2) 36, 34:5 38. 36 | 44°5, 42 | 15—25)
2-1 —1-4135-5, 34-2] 36, 35 Oe Ca i) VA eee
1-9 —1-4)/35°5, 34°2) 36, 35°2 ate 17—-26| 63
2—1°4/35°5, 34°2) 35°5, 35 44:5, 42 | 18—25
6 Fishery Board for Scotland.
INCUBATION OF HERRING Ova—continued.
|
a TEMPERATURE OF THE WATER.
A sees oy = FLow or WATER.
é (1) eevee lee
® Gravel Glass Tin cooled
= eo ie a i Plates. ; Le eae
= Range in { Max. and “Max. and Max, and | Max. and ~ Nos. | Tin ~ |Uneool’d
4 =f Min. °F. Min. °F. Min. °F. Min. °F. 1 and 2. ‘| Gravel.
| | Secs. | Secs. | Secs.
HpiD-2 1A Reames 186°5,.35:5) ee ‘oases S |) a
46 2—1°2/35°5, 34:2) 36, 35°5 at Ba eg rea ee) ee
AT 2—1 | 35:2, 34 | 36, 34:2 ne 45:5, 42 14,18 ee |
48} 18-1 35, 84 |35'5, 34°2 st 445,38) 18 ae |
49 2—1:2| 35, 34 36, 35 Pi 43°5, 42 | 16—27 ie |
50} 2:2—1:2/35- 5, 34:2) 36, 35°5 a 45°5, 44 17, 36
51! - 6°8 44, 36 44, 66 te 46, 44 9
52 es ma sh e A4°5, 43°5
53 Es Ss P =~ AT, 43
54 fa! S, ahs oe 49,47 |
oD re = | aa a 48°5, 48:5
|
* Water supply stopped for some time during the day.
RESULTS.
The Uncooled Spawn on Gravel.
This spawn was not very clean.
During incubation, the spawn was fully exposed to daylight, but
not to direct sunlight. The eggs detached from the gravel varied in
size from 1°32 to 1:5 mm. in diameter. Eggs which had been stuck
to the gravel, and which had been forcibly detached, had lost a patch
of the outer layer of the zona. This did not appear to exercise any ill
effect. No crystals were observed in this lot of spawn.
The first larva appeared on the 19th day, and larvae were
obtained almost daily up to the 29th day. On the latter date a few
of the eggs still contained embryos.
Spawn Ineubated in the Galvanized Tin.
This lot of spawn was treated with water similar to that supplied
to the uncooled gravel spawn up till the time when the larvee began
to appear, viz.,on the 20th day. Thereafter the water was cooled,
and the incubation-period was, in consequence, extended for an
additional 21 days. The larve hatched out daily from the 20th to
the 41st day. The results in this case were very satisfactory.
The retardation consisted in keeping the developed embryos in the
egg after they were ready to hatch. The result was that many of the
larvee, when they hatched, had already consumed a large portion of
the yolk. On the 40th day three had only a trace of yolk left (fig. 6).
although a fourth had still a fairly large ball of yolk unabsorbed.
A normally hatched larva may be 8 mm. in length: a post-larva
measured 7°5 or 8 mm. The eggs did not seem to have suffered from
the cooling.
The Retardation of the Development of the Ova of the Herring. 7
Diatoms were noticed at different times on the eggs, some of
which became somewhat dirty externally. They were, however,
partly sheltered from the light by being enclosed within the
pattially-closed tin. They did not, therefore, become so dirty with
diatomaceous growth as the other lots of eggs.
Crystals, in some cases in great quantity, were observed in certain
egos. The majority showed none. The crystals were attached to the
zona, sometimes in rosette-form, or even inside the embryo. They
were observed in one of the trunk canals, apparently the gut (fig. 3).
In the embryo, a large corpuscle plugged the heart with each pulsation,
and then receded again (fig. 9). Next day the corpuscle remained
clear of the heart. Crystals were observed in the gut of another
embryo, the heart of which had no plug. One lively embryo had a
huge quantity of crystals attached to the inside of the zona.
Cooled, Spawn.
The gravel spawn examined nine days after spawning seemed to be
in a pretty good condition, but some dead eggs were to be seen. On
the 11th day, the spawn on the glass plates looked well, except where
the eggs were in a thick mass. In such places dead eggs were
observed. Some of them had no doubt been killed by the pressure of
adjacent eggs. Certain eggs were of especially large diameter.
Three that contained embryos showed no crystals, but in certain of
them a granular matter was sticking on the embryo—an unhealthy
sign. An egg, 1°75 in diameter, had an unhealthy looking embryo.
Two eggs measured 1°3 and 1°35 mm. in diameter. Both contained
crystals. In one the crystals were large and few in number; in the
other they were small and fairly numerous.
By the 22nd day a copious growth of diatoms was noticed on the
eggs. The thickly-covered plates did not look so well as those that
had a sparse coating of eggs. Larve began to appear on the 29th
day. On the 42nd day a considerable quantity of dead eggs was
observed. Many appeared to the naked eye of a milky tinge. That
probably indicates that they had died recently. The milky appear-
ance is due to the perivitelline fluid turning opaque. Some of the
ova were yellow-coloured from the coating of diatoms.
On the 5lst day, ripe eggs, containing live embryos, were quite
yellowish. A good proportion of the eggs of one of the good plates
seemed to have hatched.
I examined some eggs that had died recently. In one I could
detect no movement of the embryo. I dissected it out of the egg
capsule. The heart was found to be beating slowly. The embryo
seemed to be perfect. It was, I think, dying from suffocation, due to
the mat of diatoms that covered the zona. The larve which were
obtained on each day from the 29th to the 38th day were prematurely
hatched. They were very small, viz., 4°5-6mm. long. The head was
markedly bent downwards, t.e., much more than in the older larve.
The postlarval body is shorter than normal. Succeeding batches
oradually improved in size. On the 38th day 94 were obtained.
Compared with a larva which had consumed nearly all its yolk, they
were shorter and had the head more flexed. ‘They were pretty lively.
Two measured 6 and 75mm. in length respectively. Thereafter the
8 Kishery Board for Scotland.
following quantities of fry were obtained :—39th day, 120 (20 dead)*:
40th, 250 (20 dead); 41st, 191; 42nd, 335 (100 dead); 43rd, 160
(10 dead) ; 44th, 187 (10 dead); 45th, 98 (4 dead); 46th, 67 (8 dead);
47th, 24 (2 dead) ; 48th, 27 (1 dead) ; 49th, 16 (5 dead): 50th, 14
(4 dead) ; Olst, 8 (3 dead); 52nd, 1 ; 58rd, 1 larva.
After the 39th day the larvee seemed normal. They were shorter
on the whole than the average larva, and they had a comparatively
large amount of yolk still remaining unabsorbed. The heads were
flexed. They wriggled about quite actively with eel-like movement.
On the 47th day two measured 7 and 8 mm. respectively. The head
was only slightly flexed. On the 50th day 10 good fry showed stilla
good quantity of yolk. ‘The heads were slightly flexed.
The hatching of the ova began prematurely in consequence, I
consider, of the decay of the zona, caused by the diatoms attached to
it. The same cause led, in my opinion, to the death, through
suffocation, of many ova. There was also a large quantity of infusors
and many nematodes among the eggs.
The larvee do not appear to have dev eloped much too rapidly. The
cooling was doubtless not sufficiently low. I think a slightly lower
temperature, 34° to 35° F., would not injure the ova and would
probably be sufficient to retard the ova for the requisite period.
Gravel and Glass.
While the spawn attached to the gravel did very well, that on the
glass could not be said to be distinctly inferior. Glass plates are
much more easily handled and with their aid a larger quantity of ova
can be dealt with. The spawn on the gravel was less in quantity,
less crowded, and its situation on small pieces of stone permitted,
possibly, a more effective aeration.
Kxposure to Light: Growth of Diatoms.
A danger which was apparent during this experiment has, I believe,
a greater bearing on the well-being of the ova than their location on
glass or gravel. That is, exposure to light. Light has a great
influence in stimulating the growth of diatoms as Allant and Nelson
show. If deprived of light the culture of diatoms dies off. The
quahty of the water also affects the growth of diatoms. The two
authors point out that in the tank-water of the Plymouth Laboratory
larger and healthier growths of diatoms were got than in water pro-
cured some distance off shore. This fact was ascribed to the greater
quantity of organic matter in the tank-water.
The diatoms which grow on the ova tend to prevent the aeration of
the embryo, and are also, I consider, the probable cause of the
premature escape of the larve by hastening the decay of the zona.
No doubt decay takes place normally during incubation. If that
decay be hastened the embryo may be able to burst its way to
freedom at a smaller size than the av erage.
* The larvee were caught in a trap in which some were killed.
+ ‘The Artificial Culture of Marine Plankton Organisms.” Quarterly
Journal of Microscopical Science, Vol. 55, Pl. 2. June, 1910.
Diatoms attached to the eggs of Lobster, exposed to bright light, caused the
decay of the outer egg-membrane. (Anderton) Report of “Marine Department,
New Zealand, 1908-9. Wellington, 1909.
The Retardation of the Development of the Ova of the Herring. 9
The Current of Water.
I think that the current of water should be strong. It is not
necessary that it be continuous: it may be intermittent. A large
quantity sent through the apparatus at intervals would probably be
more effective than a small continuous current.
The Filter-Barrel.
During the experiment, the filter-barrel, in which was filtered the
water that was cooled, was allowed to work for 40 days. This was
probably an error. Whether it affected the success of the experiment
is donbtful. It would be better to have the filters frequently cleaned.
EXPERIMENTS WITH SPAWN FROM LIVE AND DEAD HERRINGS.
On Galvanized Wire Gauze and Glass Plates.
A number of ripe herrings were put into a herring barrel and sent
to Aberdeen. On arrival all but two were dead. The survivors were
a male and a female. The herrings had been probably not more than
12 hours in the barrel. Four experiments were carried out on March
24th 1911.
(A) Some spawn was obtained from the two live fishes and it was
put on a piece of galvanized wire gauze. The gauze was new, but it
had been for a day or two in running sea-water. The eggs appeared
to do well for a few days. They showed lots of crystals. The eggs
gradually died off, and on the 25th day after fertilization all but three
were dead. The embryo at that time showed black pigment in the
eyes.
* (B) Spawn and milt obtained from dead herrings were put on
narrow-meshed galvanized wire gauze in hatchery water. The eggs
developed for a time, but by the 20th day all but one or two were dead.
(C) Spawn was obtained from a dead fish and put on a glass plate.
This spawn was put into the water in which the herring travelled to
Aberdeen. No fresh milt was supplied to the water. I examined a
drop of the water: tailed sperms were visible, but they showed little
motion. Fertilization ensued, however. Although most of the eggs
died off, two which had embryos ready to hatch were found alive on
the 31st day. A good number of the eggs had died after pigment
had began to show in the eyes of the embryo.
(D) Some spawn and milt were taken from dead herrings and put
on a glass plate in new (hatchery) water. Fertilization took place in
this case also. Many of the eggs had well developed embryos, but
most died before hatching.
The four lots of spawn stood, after being prepared, for some hours
in still water. They were then all put in a revolving tin box set in a
galvanized iron tank. The water supplied to them was cooled. ‘The
temperature is given in the column marked “Tin” (p. 5) from the date
March 26th onwards. One egg which had been in still water all night
showed a large normally segmented disc and a huge quantity of
crystals. Some fry were got from the spawn on the 26th to 31 days
after fertilization.
10 Fishery Board for Scotland.
Milt and Spawn.
Some milt from a dead herring was put into sea-water. I found the
sperms (fig. 7) oscillating vigorously as if anchored by the tail.
Many, however, oscillated very faintly.
No difference was observed between the spawn of the living and
dead herrings.
Some unfertilized eggs, a quarter of an hour after they had been
deposited on the glass plate, exhibited different shapes (fig. 8). Most
of them show an inpushing where they had been pressed by another
egg. No sign of perivitelline space was visible. The eggs were
easily dislodged from the glass. They do not adhere strongly until the
perivitelline space is for med, Three days later these unfertilized eggs
had a fairly large perivitelline space, but they remained dull tinted on
the outside, instead of showing a clear translucent zona.
THE CRYSTALS IN THE EGGS.
The crystals, which were very evident during the two previous
experiments, were found also on this occasion. They are rectangular
(fig. 5), sometimes ending in a tapering oblique point. They may be
fine, needle-like, or fairly thick bars. They appeared in the batches
of eggs which were exposed to the influence of galvanized iron. In
one instance, they appeared within 56 hours after fertilization (fig. 10).
This ege had been in still water in an enamelled bath for a night, and
had probably with it some galvanized gauze.
No crystals were observed in the uncooled eggs on gravel. The
water was delivered to these eggs through large g galvanized iron pipes,
which had been in use for several years. All the cooled spawn
received water through a series of small galvanized iron pipes.
One remarkable fact is that the crystals were not present in all the
eggs of a batch. They may be found (1) on the inside of the zona ;
(2) between the layers of the zona; (5) attached to the yolk or to the
embryo; and (4) within the embryo.
These crystals are soluble in acetic acid, and in sodium hydrate
solution. They are insoluble in water, alcohol, and methvlic ether.
Dr. J. K. Wood, Chemical Department, ‘University College, Dundee,
says that the er ystals, being able to act both as a base and an acid, are
probably formed from proteids by some process of decomposition.
I have came to regard the galvanized iron as, in some measure, the
cause of the formation of these crystals. The galvanized iron becomes
coated with a white incrustation in sea-water. Dr. Wood was not
able to find zinc in solution in the sea-water in which a piece of this
metal had been lying for several days. It seems possible that the
cause of the formation of the crystals is to be found in electrolytic
action. The electric current acting within the eggs on the fluids con-
taining excretory products causes the crystallization. I formerly
regarded the crystals as indicative of deficient aeration, and [I still
favour that view. The deficient aeration resulting in the accumulation
of excretory products in the perivitelline fluid may afford the oppor-
tunity for the action of the current. All the eggs do not exhibit the
crystals. It is possible that the zona may not be acting satisfactorily
from the point of view of osmosis. It is possible that variation may
The Retardation of the Development of the Ova of the Herring. 11
occur in that capsule. As I pointed out above, the diameter of the
zona varies much. In some eggs the two layers of the zona may be
seen separated. In one egg here and there the vitelline membrane
may be made out. Further, the eggs are differently served by the
currents of water.
It is quite possible that the quality of the water at the Laboratory
may be a contributory factor. It is probably charged with organic
matter.
While active embryos are found in eggs containing the crystals, it
is not likely that their presence is an advantage. When the crystals
are formed in a canal in the embryo, they will almost surely prove a
fatal encumbrance.
RECOMMENDATIONS.
If the experiment of transporting the ova of the herring to New
Zealand be made, the following would, I think, be a suitable appa-
ratus. It is shown in plan in fig. 1. It consists of six compartments,
each of which contains a revolving frame. The frame will be able to
carry on its external faces four glass plates, 9 inches by 7 inches. A
section through two of the compartments is given in fig. 1a. A shaft
running longitudinally above the middle of the apparatus would
impart the revolving motion to the frames by means of little belts
(rubber cord.)
The amount of water circulated through the apparatus should be as
large as possible. If iron pipes are used to cool the water it will
probably be better to have ordinary unprotected iron pipes.* In any
case, the pipes and apparatus should be well seasoned, by being
exposed to the action of sea-water, for some time previous to the
experiment being made. ‘The water when it leaves the cooling
apparatus should pass into a reservoir, surrounded by ice. Thence it
should flow through the apparatus. If the water be not used over
again, the waste water can be utilized to cool the mcoming water.
Where the water is to be used over again, a pump will be required to
bring it from the reservoir. A small quantity of freshly-cooled water
should be steadily added.
This apparatus should be provided with lids, and should be kept in
a dimly-lghted apartment.
It is most important to have clean water at all times for the spawn.
This should be carefully arranged when the spawning is taking place,
for mud, etc., adhere readily to the ova. “Wherever necessary, the
water should be filtered through sand. It is hardly possible that all
the eggs pressed on to the plates will live.
Where only a small quantity of cooled water is available, it might
be advisable to have it sterilized. I have not, however, made any
experiments with sterilized water.
Some precautions should be observed when the spawn is being
obtained. Only spawn that runs freely on gentle pressure should be
employed. Milt and spawn should be preferably obtained from living
fishes. The eggs should be arranged thinly on the plate. They
should be protected from being touched after they are on the plate.
The spawn should be brought into moving water as soon as possible.
_ * Vide Davis., ‘‘The Action of Water on Zine and Galvanized Tron.” The
Journal of Chemical Industry. Vol. XVTII., Feb. 28, 1899.
12 Fishery Board for Scotland.
An interval of four to six hours in quiescent water has, however,
usually been given to allow of the ova becoming fastened to the glass.
The glass plates can be put into wooden boxes, each holding
about six plates. The boxes may be put in a herring barrel into a
frame, which can be revolved from time to time. The barrel would,
if necessary, be surrounded with ice during the railway journey.
The spawn should be protected from the light.
For the woodwork, I think coal-tar is a suitable preservative.
PEATE: 1
Fic. 1. Drawing (plan) of apparatus for conveyance of herring Spahite
ae A: Be (section) »
» 2. Drawing of apparatus in which the cooled herring | spawn was kept
during the Spring of 1911.
Enlarged drawing of part of embyro, to show crystals in gut.
Live egg attached to dead egg, showing large number of crystals.
Enlarged drawing of crystal.
Enlarged drawing of anterior end of a larva, which had been retained in
the egg until much of the yolk had been absorbed.
7. Enlarged drawing of a sperm.
» 8. Different shapes assumed by unfertilized eggs when they fell on the
glass plate.
,, 9. Drawing to show the corpuscle which alternately blocked and receded
from the heart of the embryo partially shown in fig. 3.
,, LO. Egg in which erystals had appeared within 36 hours of spawning.
LETTERS USED.
a.—Anus. nt.—Notochord.
er.—Crystal. o.—Outer skin of zona.
de.— Dead egg, pl.—€ plate.
dmf.—Dorsal marginal fin. t.—Tray.
gt.—Gut. | vmf.—Ventral marginal fin.
le.—Live egg. yk.—Yolk.
f ice
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