262
E.
F.
G.
H.
I.
J.
K.
L.
M.
N.
0.
P.
Q.
R.
S.
T.
U.
V.
W.
x.
Y.
2.
J. H. 5. BLAXTER AXD F. 0. T. HOLLIDAY
.. .. .. .. .. ..
Growth .. .. ..
Natural Mortality
. . .. .. .. .. . . ..
Composition of the Tisaues . . .. .. .. .. ..
Salinity Tolerance and Osmo-regulation
. . .. .. . .
Temperature, Oxygen, CO, and H,S aa Limiting Factors. .
..
The Brain
. . .. .. .. .. . . . . ..
Vision . . .. .. .. .. .. .. .. ..
Olfaction. . .. ..
The Labyrinth, Hearing, theEffeot of Sound, and Sound Production
. . .. .. .. .. .. .. ..
Buoyancy and Equilibrium . . .. .. .. . . ..
swimming
. . .. .. .. .. .. .. ..
Activity . . .. .. .. .. .. .. .. ..
Shoaling . . .. .. .. .. .. .. .. ..
Migrations
. . .. .. .. * . . . .. ..
Vertical Migration
. . .. .. .. .. .. ..
Effect of the Moon
. . . . . . . . .. * . ..
Attraction to Artificial Light8
.. .. . . .. ..
Reaction to Nets and Other Obstacles
. .
.. .. ..
Learning. . .. .. .. .. .. ,. .. . .
Maturation of the Gonads
. . .. .. .. .. ..
. . .. .. .. . . ..
302
305
307
309
312
313
314
316
316
320
322
325
326
332
342
347
36 1
356
366
348
Spawning
.. .. .. . . .. .. . . . . 364
Racial Characters, the Genotype and the Environment . . .. 361
VII. Conclusions
. . .. . . .. .. .. .. . . .. 370
VIII. References
. . .. .. .. .. .. .. .. .. 372
I. INTRODUCTION
A. General
Of all the clupeids the herring, Clupea harengw, has been most
extensively studied. Perhaps it has been more the subject of research
than any other fish species, partly due to its commercial importance and
partly to the need for so much work before even tho ~ ~ s c r i t i a l ~
of i t H
complex biology could be exposed ; tliw the herring i H bound to take u p
tho main H ~ C O
in this review. Although most of the retlearch has been
conconic~d with the racial question and population studies, these aspects
are only mentioncd where experimental work has assisted such research.
Therefore while natural mortality due to disease and predation, and
fecundity in relation to reproductive biology, will be considered,
fishing mortality and its estimation, and fecundity as a means of
distinguishing races and for estimating stock size, will not.
For a general review of clupeid biology reference may be made to a
recent bibliography (Scattergood, 1957), to a series of translations from
Russian (Scattergood, 1959), to the monograph on tho Clupeidac by
Svetovidov (1952) and to general books Ruch aa those of Hodgson (1067)
and Nikolsky (1957). Two recent Hpecial scientific meetirign of t h
International Council for t h o 12xploration of tho Soa in (hpnhngun
(J.C.N.S.) in 1 f W I and 1901 c:ovcmrtl much huhaviour and physiological
E.
F.
G.
H.
I.
J.
K.
L.
M.
N.
0.
P.
Q.
R.
S.
T.
U.
V.
W.
x.
Y.
2.
J. H. 5. BLAXTER AXD F. 0. T. HOLLIDAY
.. .. .. .. .. ..
Growth .. .. ..
Natural Mortality
. . .. .. .. .. . . ..
Composition of the Tisaues . . .. .. .. .. ..
Salinity Tolerance and Osmo-regulation
. . .. .. . .
Temperature, Oxygen, CO, and H,S aa Limiting Factors. .
..
The Brain
. . .. .. .. .. . . . . ..
Vision . . .. .. .. .. .. .. .. ..
Olfaction. . .. ..
The Labyrinth, Hearing, theEffeot of Sound, and Sound Production
. . .. .. .. .. .. .. ..
Buoyancy and Equilibrium . . .. .. .. . . ..
swimming
. . .. .. .. .. .. .. ..
Activity . . .. .. .. .. .. .. .. ..
Shoaling . . .. .. .. .. .. .. .. ..
Migrations
. . .. .. .. * . . . .. ..
Vertical Migration
. . .. .. .. .. .. ..
Effect of the Moon
. . . . . . . . .. * . ..
Attraction to Artificial Light8
.. .. . . .. ..
Reaction to Nets and Other Obstacles
. .
.. .. ..
Learning. . .. .. .. .. .. ,. .. . .
Maturation of the Gonads
. . .. .. .. .. ..
. . .. .. .. . . ..
302
305
307
309
312
313
314
316
316
320
322
325
326
332
342
347
36 1
356
366
348
Spawning
.. .. .. . . .. .. . . . . 364
Racial Characters, the Genotype and the Environment . . .. 361
VII. Conclusions
. . .. . . .. .. .. .. . . .. 370
VIII. References
. . .. .. .. .. .. .. .. .. 372
I. INTRODUCTION
A. General
Of all the clupeids the herring, Clupea harengw, has been most
extensively studied. Perhaps it has been more the subject of research
than any other fish species, partly due to its commercial importance and
partly to the need for so much work before even tho ~ ~ s c r i t i a l ~
of i t H
complex biology could be exposed ; tliw the herring i H bound to take u p
tho main H ~ C O
in this review. Although most of the retlearch has been
conconic~d with the racial question and population studies, these aspects
are only mentioncd where experimental work has assisted such research.
Therefore while natural mortality due to disease and predation, and
fecundity in relation to reproductive biology, will be considered,
fishing mortality and its estimation, and fecundity as a means of
distinguishing races and for estimating stock size, will not.
For a general review of clupeid biology reference may be made to a
recent bibliography (Scattergood, 1957), to a series of translations from
Russian (Scattergood, 1959), to the monograph on tho Clupeidac by
Svetovidov (1952) and to general books Ruch aa those of Hodgson (1067)
and Nikolsky (1957). Two recent Hpecial scientific meetirign of t h
International Council for t h o 12xploration of tho Soa in (hpnhngun
(J.C.N.S.) in 1 f W I and 1901 c:ovcmrtl much huhaviour and physiological