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FISHERIES AND MARINE SERVICE
Translation Series No. 3831
Gonadotropic and sex hormones and their role in regulating
the reproductive system in cold-blooded vertebrates
by I.A. Barannikova
Gonadotropnye i polovye gormony i ikh rol' v
Original title:
regulyatsii funktsii vosproizvodititel' noi sistemy u
poikilotermnykh pozvonochnykh
From.,
Tr. Vses. Nauchno-Issled. Inst. Morsk. Rybn. Khoz. Okeanogr.
111: 34-53, 1975
Translated by the Translation Bureau( JW
Multilingual Services Division
Department of the Secretary of State of Canada
Department of the Environment
Fisheries and Marine Service
Vancouver Laboratory
Vancouver, B.C.
1976
37
pages typescript
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AUTHOR - AUTEUR
I.A. t3arannikova
TITLE IN ENGLISH - TITRE ANGLAIS
Gonadotropic and sex hormones and their role in regulating the
reproductive system in cold-blooded vertebrates
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Gonadotropnye i polovye gormony i ikh rol' v regulyatsii funktsii
vosproizvodititaJ6noi sistemy u poikilotermnykh pozvonochnykh
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From: VNIRO (All-Union Research Institute of Sea Fisheries and Oceanography),
vol. 111, 1975, pp. 34-53 (USSR). Trudy.
Gonadotropic and sex hormones and their role
in regulating the reproductive system in
UNEDITED TRANSLATIOle
For information only
cold-blooded vertebrates
by
TRADUCTION NON REVISLI
Information seulernont
I.A. Bàrannikova
UDC 597--114:597.442
The reproductive system in cold-blooded vertebrates is regulated by
various components of the neurocrine and endocrine systems. These systems,
compared to those in the Higher vertebrates, have a peculiar structure and
function and their hormons interact in characteristic ways in regulating the
reproductive system. These topics are discussed in a number of papers of
the present collection [Barannikova, Polenov] and hence will not be fully
reviewed here.
The main focus will fall on the role of the gonadotropic
and sex
hormones in the reproductive system of fish. A short discussion of the same
problems with regard to the ancient Agnatha vertebrates and Higher classes
of cold-blooded vertebrates such as amphibians and reptiles, in comparison
with fishes, makes it possible to regard these problems from an evolutionary
point of view.
*The numerals in the right-hand margin indicate page numbers of the
original (Tr).
SEC 5-25T (6/76)
34*
2
In cold-blooded vertebrates the gonadotropic hormones of the pituitary play a dominant role in regulating gonad functions.
The gonads of the
Lower vertebrates secrete a whole series of sex hormones (androgens and
estrogens) that interact in complex ways with gonadotropic hormones and play
a large role in the reproduction process (Fontaine, 1969; Fontaine, 1972;
Hoar, 1965, 1969).
Among the factors that regulate the reproductive system in most coldblooded vertebrates, the seasonal and cyclical functions of the gonads and
pituitary gland are extremely important. On the basis of a study of sex
cycles and comparisons of the condition of sex glands with functional changes
in the pituitary at different periods, we have obtained data on the gonadotropic function of the pituitary gland in fish and on the cells that secrete
a gonadotropic hormone (Gerbirsky, 1940, 1947; Barannikova, 1949; Pickford,
Atz, 1957; Barr, Hobson, 1964, and others).
Along with this type of study for determining the role of the gonadotropic and sex hormones, important experimental methods, such as removal of
the pituitary gland and gonads, were also used at various times of the life
cycle with subsequent suitable therapy.
The effect of hormones of various
vertebrates (from fishes to mammals) was investigated, which makes it possible
to examine this problem with regard to the phylogenetic specificity of the
hormones connected with reproduction.
In contrast to the well-known papers dealing with the physiology of
the pituitary and sex glands in lower vertebrates (Pickford, Atz, 1957; Dodd,
1960; Hoar, 1955, and others), our paper discusses mainly recent data.
3
Y
Gonadotropic hormones
In the Higher vertebrates the pituitary secretes two gonadotropic
hormones -- a follicle-stimulating hormone (FSH), which induces growth and
maturation of the ovarian follicle in females and spermatogenesis in males,
and a luteinizing hormone (LH), which stimulates the secretion of estrogens
and ovulation in females and affects the interstitial cells of the testis
and induces the sythesis of androgens. The luteotropic hormone, or prolactin
(LTH), is also associated with gonadotropic activity.
As to the Lower vertebrates, at present there are no precise data
regarding the presence in their pituitary gland of the two different gonadotropic hormones.
This question is under intensive study and general infor-
mation on this point has appeared in a number of papers (Pickford, Atz, 1957;
Pickford, 1959; Atz, Pickford, 1964; Bern, Nandi, 1964; Barr, 1965; Hoar,
1965, 1966, 1969; Matty, 1966; Dodd, Wiebe, 1968; Ball, Baker, 1969; Sage,
Bern, 1971; Turdakov, 1972; Barannikova, 1973; Danaldson, 1973; Schreibman
et al., 1973).
Interesting data have been obtained on the purification of fish
gonadotropins, involving mostly carp (Burzawa-Gerard 1971) and chinook salmon
(Donaldson et al., 1972; Donaldson, 1973).
These studies showed that the molecular weight of the gonadotropin
of carp and chinook salmon falls within 27,000-30,000, but there is a subunit with a smaller molecular weight.
A study of the chemical properties of
fish gonadotropins has shown that by its amino acid composition this preparation is closer to the FSH than to the LH of mammals. However, at present the
chemical characteristics of gonadotropins of Lower vertebrates are not very
well known, and to investigate them is one of the essential tasks of comparative and evolutionary endocrinology.
4
Cyclostomes. In river lamprey (Lampetra fluviatilis) removal of the pituitary gland resulted in inhibited growth of the ovary and testis.
In males
spermatogenesis was inhibited, but the process continued if some spermatogonia
had already divided by
the time the pituitary gland was removed.
In female
lampreys removal of the pituitary gland does not result in atresia of the
ovarian follicles, which occurs in all other vertebrates (Dodd, 1960).
Removal of the pro- and meso-adenohypophysis (homolog pars distalis) allowed
the vitellogenesis in lamprey to continue, but ovulation did not occur if the
operation was made in the fall or winter (Larsen, 1965, 1969a, b). If the
hypophysectomy was made after the follicular cells had begun to grow (spring),
the oocytes continued to grow and ovulation occurred in some lampreys. In
males, spermiation occurred if the pituitary gland was removed in the spring
(Larsen 1973).
Hence in cyclostomes gametogenesis can continue in the absence of
gonadotropic hormones, which distinguishes this group from other vertebrates
in which removal of the pituitary precludes the occurrence of meiosis and the
conversion of spermatogonia into spermatocytes (in males) (Evennett, Dodd,
1963; Wiebe, 1968). This suggests that in this ancient group of vertebrates
(Agnatha) the development of the sex cells occurs to a significant extent
independently of the pituitary hormones.
It has been demonstrated that the pituitary gland of the lamprey
induces gonad maturation in animals of the same species (Lanzing, 1959, Even-
/36
nett, Dodd, 1963) and that intact lampreys react favorably to carp pituitary
preparations (Lebkova, 1956) and to mammalian gonadotropic hormones (Dodd,
1960; Evennett, Dodd, 1963). In hypophysectomized lampreys there appeared
no secondary sexual characters whose development is regulated by steroid
hormones.
This suggest that the lamprey gonads contain endocrine tissue which
is regulated by the pituitary gland (Pickford, Atz, 1957; Dodd, 1960; Larsen, 1973).
Thus the gonadotropic and steroid hormones participate in the regulation of the reproductive system of the ancient members of Agnatha vertebrates.
However, the interrelationship between the pituitary and the gonads
in lampreys is quite distinctive. Lampreys are the only vertebrates in which
the sex cells grow and mature to a significant degree independently of the
influence of gonadotropins and hence there is no precise reciprocal relationship (at certain periods, in any case) between the pituitary gland and the
gonads.
In this connection, it should be noted that in lampreys ovulation
can occur by auto transplantation of the pituitary substances outside of any
links with the brain (Larsen, 1973), which suggests that the hypothalamic
regulation of this gland in the cyclostomes differs from that of other vertebrates.
Elasmobranchs.
Experiments in removing the pituitary gland from selachian
fishes are very revealing (Dodd, 1955, 1960; Pickford, Atz, 1957 --combined
data from previous papers).
The pituitary gland of these fishes has precise segments which can
be removed separately, and the effect on the gonads can then be studied.
When one segment-- the ventral portion of the pituitary body--was removed
from the shark Scyliorhinus caniculus, it resulted in follicular atresia and
stopped the vitellogenesis in the oocytes.
Removal of the entire pituitary
gland has similar results (Hisaw, 1959; Chieffi, 1962). _
The absence of gonadotropins in the testis makes it impossible for
the spermatogonia to divide and to form spermatocytes and thus the later phases
of sex cell growth in males degenerate. This has been demonstrated in several species of sharks (Dodd et al., 1960; Simpson, Wardle, 1967).
.
6
Thus experiments with elasmobranch fishes have provided convincing
evidence that certain gametogenic phases are regulated by the gonadotropic
hormones (Dodd, Wiebe, 1968).
It is likely that the pituitary gland in
elasmobranchs, as in the Higher vertebrates, regulates.the steroid-producing
endocrine tissue of the gonads.
The gonadotropic activity of the pituitary
varies during the reproductive cycle (Dodd, 1972).
These data present convincing evidence that the cells that generate
gonadotropins in elasmobranch fishes are located in the ventral part of the
pituitary body.
However, the gonadotropic elements of these fishes have not
been identified in detail (Ball, Backer, 1969).
The gonadotropins of elasmobranch fishes have not been investigated
biochemically, but the presence of gonadotropins similar to FSH and LH is
assumed, based on ordinary tests with rats (Witschi, 1955).
Teleosts. The pituitary gland of these fishes has been described, based on
ecological histophysiology (Gerbil'sky, 1947, 1956, 1960) and experimental
analysis, as containing basophilic (cyanophilic) cells that generate a gonadotropic hormone.
The histophysiology of the pituitary gland of teleosts has
been studied in greatest detail, whereas the cartilaginous ganoids and particularly the bony ganoids are less well-known (Gerbil'sky, 1940, 1947;
Pickford, Atz, 1957; Barannikova, 1949, 1969; Moiseeva, 1969, 1970, 1971;
Barr, Hobson, 1964, and others).
The largest hormone quantity that induces
ovulation, movement of the sperm from the ampullae into the discharge ducts
of the testis and transformsthe fish organism into spawning condition is contained in the pituitary shortly before spawning. These data formed the basis
for developing the method of hypophyseal injections which are so widely used
in fish management (Gerbil'sky, 1941, 1947).
/37
7
At present the main problem in the study of the gonadotropic function
of the pituitary has to do with determining the number of hormones this gland
secretes.
Along with solving this problem, a study of the cytophysiology of
the pituitary is very important. Histochemical and electron-microscopic
methods have enabled us to identify only one type of gonadotropic cells in
a number of fish species (Gasterosteus aculeatus, Follenius, 1968; Leatherland, 1970; Cymatogaster aggregate, Leatherland, 1969, and others).
But
there is considerably more information available on the existence of two types
of gonadotropic cells, differeing in form, size, type of inclusions and degree
of functional activity at various gametogenic stages. Information on these
questions has appeared in a series of papers (Ball, Baker, 1969; Hoar, 1969;
Sage, Bern, 1971; Schreibman et al., 1973; Donaldson, 1973).
A study of the ultrastructure of the pituitary in sockeye salmon
during the maturation period has produced valuable data.
Several studies
of the posterior lobe of the pituitary of this species, which were conducted
with a light microscope, revealed only one type of gonadotropic cells (van
Overbeeke, McBride, 1967).
A study using electron microscopy revealed the
presence of two types of gonadotropic cells: one of them contains small,
electron-dense granules, and the other contains larger, less dense granules.
These different types of cells also differ according to periods of maximum
sexual activity (Cook, van Overbeeke, 1972).
However, the presence of two
types of gonadotropic elements in the pituitary of a number of teleost species
does not prove that these fishes have two gonadotropic hormones.
The use of immunochemical methods has made it possible to indentify,
in the pituitary of sockeye salmon and carp, cells which together with an
antiserum react to an LH and which consequently are gonadotropic elements
(McKeown, van Overbeeke, 1971; Billard et al., 1971).
8
The method of immunodiffusion has revealed changes in the antigen
content of the acetonated pituitary of Gobius melanostomus at various stages
of the sex cycle; it has revealed and quantitatively assessed the antigen
factors connected with the gonadotropic function of the pituitary of this
species.
An examination of the antigen factor connected with the gonado-
tropic function of the pituitary revealed a dissimilarity in the precipitate:
it is most often split. This is important in connection with the question
regarding the number of gonadotropins in the pituitary of Gobius melanostomus
(Apekin, Moiseeva, 1973).
The experiments in hypophysectomy and-the subsequent therapy with
fish pituitary preparations and gonadotropins of Higher vertebrates played
an important role in determining the number of gonadotropic hormones and
their effects in fish.
Hypophysectomy was performed on over 20 species of teleosts (Pickford,
Atz, 1957--contains references to earlier papers on this subject; Barr, 1963a,
b; Yamazaki, 1965; Ahsan, 1966; Liley, 1969; Pandey, 1969; Hoar, 1969, and
others) and on cartilaginous ganoids (sturgeons) (Zubova, 1969). None of
the hypophysectomized fish revealed any spermatogonial division into spermatocytes.
A reduction in mitotic activity of the spermatogonia was observed
in Couesius plumbeus.
In this species spermatogenesis resumed when a salmon
pituitary preparation or a mammalian LH was administered,_whereas a folliclestimulating hormone remained ineffective (Ahsan, 1966). Similar results were
obtained with Fundulus heteroclitus (Lofts et al., 1966).
Administration of
a LH in combination with a somatotropic hormone induced spermatogonic mitosis
and further spermatogenesis.
It is interesting that the main effect of admin-
istering LH to hypophysectomized males of this species was that it stimulated
the interstitial cells and increased the activity of 3S-hydroxysteroid dehydrogenase (3G IISD), which suggests increased steroid synthesis.
Asa result, the
/38
hypophysectomized specimens developed spermatozoids and spermiation (at a
rather high temperature) (Pickford et al., 1972). In gold fish spermiation
is caused by the effect of gonadotropins on steroid synthesis (Yamazaki,
Donaldson, 1969).
Hypophysectomy in guppies has shown that this operation causes
regressive changes in Sertoli cells and in the interstitial tissue of the
gonads (Pandey, 1969).
In females of the genus Poecilia the spawning behaviour
that was interrupted by hypophysectomy can be restored by administering a
Salmon pituitary preparation thus affecting the steroid tissue of the gonads
(Liley, Donaldson, 1969).
These data corroborate the notion that the steroid
tissue of the gonads is regulated by gonadotropic hormones of the pituitary
gland.
Thus the gonadotropic substances of the pituitary have a significant
effect on the production of steroids in the sex glands. The sex hormones, in
turn, determine the growth of secondary sex characteristics and bring about
certain growth and maturation phases of the sex cells.
In hypophysectomized females the protoplasmic oocytes remain undamaged, even though mitotic division of the oogonium ceases (Yamazaki, 1965).
Oogonial mitosis can be regenerated by administering fish pituitary preparations.
Hypophysectomy always results in inhibited vitellogenesis and atrophy
of oocytes containing a yolk. LH has a slight stimulating effect on oocyte
growth in the sticklebacks (Ahsan, Hoar, 1963). However, not all fish showed
a similar effect.
Administration of mammalian gonadotropins did not induce
yolk growth in hypophysectomized gold fish (Yamazaki, 1965).
At present, a number of fish species have been studied with the use
of methallibure--a substance that blocks the synthesis of gonadotropins in
the pituitary without affecting the neurosecretion of the hypothalamus.
10
Administration of this preparation, as has been shown on the basis of
Cymatogaster aggregata, Carassius auratus, Gasterosteus aculeatus, results
in suppression of spermatogonial mitosis, termination of vitellogenesis and
spermatogenesis and in reduced production of steroid hormones.
Administra-
tion of a carp pituitary preparation and mammalian LH induced renewed spermatogenesis and triggered secretory activity of the interstitial cells.
The
FSH again had no effect (diebe, 1968). FSH also had no effect in hypophysectomized small catfishes (Sundararaj, Nayyar, 1967). When young guppies are
exposed to methallibure, only spermatogonia can be detected in the testis
(Pandey, 1970; Leatherland, 1970).
The same dosage in Tilapia prevented spermatogonial division into
spermatocytes and sperm secretion, reduced the number and size of interstitial
cells and the amount of testosterone in the testis.
oocytes that contained a yolk underwent atresia.
In the females, all
Administration of Tilapia
pituitary preparations to male and female Tilapia neutralized most of the
effect of methallibure. In males, the effect of methallibure was also largely
neutralized by administration of chorion, but the latter had no effect in
females, whereas administration of FSH restored normal ovary function.
This
suggested to the investigators that the Tilapia pituitary gland secretes two
gonadotropic hormones, or one gonadotropin with two different subunits (Hyder,
1972).
Adding methallibure to the food of Tilapia mossambica and T. aurea
not only affected the sex cells of these fish but resulted in the extinction
of secondary sexual characteristics and spawning
behaviour.
Also, the meso-
adenohypophysis in these fish contained fewer basophilic cells and less
intensive coloration (Dadzie, 1972).
139
11
Hence in fish males the gonadotropic hormone induces spermatogonial
mitosis and is required for the division of spermatogonia into spermatocytes,
and in females the gonadotropic hormone is required for oogonial mitosis,
vitellogenesis and further maturation of the egg cells. The stimulating
effect of the gonadotropic hormone on the generation of steroids in the sex
cells in males and females has been well demonstrated. A similar relation
has also been established for the elasmobranchs (see above).
Very interesting data have been obtained on the part played by LH
in regenerating spermatogenesis in hypophysectomized fish.
We know that in
mammals FSH regulates the spermatogenic activity of germinal epithelium,
whereas LH stimulates the secretory activity of the interstitial tissue. In
fish, as these data indicate, FSH has no stimulating effect on the testis of
hypophysectomized specimens, whereas mammalian LH regenerates spermatogenesis
and activates the interstitial tissue.
Administration of a fish pituitary preparation to intact specimens
in various stages of maturation made it possible to identify the gametogenetic
moments when the effect of the gonadotropic hormone is most evident. The
effect of acetonated carp pituitary preparations (obtained from specimens
whose gonads were in the 4th stage of maturation) on ruffe (Acerina cernua
L.) resulted in stronger oogonial mitosis (Travkina, 1971) and in stronger
vitellogenesis and higher follicle cells in whitefish (Coregonus lavaretus
pidschian Gmelin) and in ruffe (Sakun, 1970; Travkina, 1972).
Adminstration
of small doses of the same preparation to ruffe induced vitellogenesis in
protoplasmic oocytes (Travkina, 1972).
A similar effect and heightened vitel-
logenesis were obtained in Tilapia (Tilapia mossambica Peters) by adminstration
of acetonated pituitary preparations obtained from humpback salmon in the 4th
and 5th stages of maturation and by administration of pituitary preparations
obtained from female Tilapia (Chistova, 1971, 1973).
12
Meiosis and ovulation in male and female fish with sex glands in the
4th stage of maturation can be obtained by administering the same hormone
preparation that was used for inducing vitellogenesis in a number of preceding
experiments. All these mitotic effects--the change of the oocytes from the
protoplasmic to the trophoplasmic stage and the heightened vitellogenesis
as well as meiosis and ovulation--were obtained by administering, to intact
specimens, acetonated pituitary preparations of fish (carp and humpback
salmon) with sex glands in the 4th stage of maturation.
Prolonged administration, to inunature humpback males, of a partly
purified gonadotropin obtained from chinook salmon (close to maturity)
stimulated spermatogonial mitosis and accelerated the spermatogenetic process.
The interstitial cells which synthesized the steroids became activated at
the same time. As a result, the males developed mature sperm one year earlier
than would have been the case under normal conditions (Funk, Donaldson, 1972).
Since in these experiments the salmon gonadotropin stimulated all phases of
the sex cells, from spermatogonial division to maturation, it is assumed that
these fish have only one gonadotropic hormone with a multipurpose effect
(Donaldson et al., 1972).
The effect of gonadotropins on meiosis and ovulation in fish (cartilaginous ganoids, bony fishes) has been studied in particular detail, including the morphology and dynamics of meiosis in vivo and in vitro under the
influence of gonadotropins (Kazansky, 1954; Detlaf, 1960, 1966; Belyaeva,
Cherfas, 1965).
It has been clearly demonstrated that the gonadotropic
hormone affects the cells of the follicular epithelium (sturgeons), resulting
in the production in these cells of a substance similar to progesterone which
causes the egg cells to mature (Detlaf and others, 1968).
S
13
Regardless whether the fish pituitary secretes one or two gonadotropic hormones, a discussion of the subject should include an examination
of the effect of various mammalian gonadotropins (FSH and LH, chorion) on
the sex glands of fish. This information is also interesting from the point
of view of the phylogenetic specificity of gonadotropic hormones which have
already been partly discussed (Pickford, Atz, 1957; Barr, 1968; Barannikova,
1969; Barannikova and others, present collection).
The attempts to use mammalian gonadotropins to induce ovulation in
fish yielded no positive' effects for FSH.
An exception in that regard is
the work of Ramaswami and Lakshman (1958). However, it is not certain whether
or not the FSH preparation they used was sufficiently pure. The use of LH
has provided positive results for a number of fish species (Ramaswami,
Lakshman, 1958; Ball, 1960; Dodd, 1960; Ramaswami, Hoar, 1966). The LH has
a rather wide-ranging effect on fish, outnumbering those effects this substance
has on mammals. These data were discussed in part above. The use of a
chorionic gonadotropin also induces ovulation in a number of fish species,
although a positive effect was not obtained in all the fish involved in the
study (Ramaswami, Sundararaj, 1957; Sneed, Clemens, 1959; Gerbil'sky, 1964;
Barannikova and others, 1969 and present collection).
Spermiation in amphibians is considered a specific test for LH. This
test confirms the LH-like activity of the pituitary in fish and amphibians
(Barr, Hobson, 1964; Hoar, 1966, and others). But, at the same time, the
reactions that are usually induced by FSH can be initiated by administering
a
fish pituitary hormones.
Testing unpurified (Kirschenblat, 1949; Witschi, 1955) and partly
purified (Otsuka, 1956a, b) fish pituitary hormones on mammals resulted in
identifying two gonadotropic hormones.
However, it is doubtful whether the
mammal test specimens can be used for identifying the number of gonadotropins
14
in the pituitary of fish (Woodhead, 1961; Burzawa-Gerard, Fontaine, 1965).
Extensive testing of purified gonadotropins of carp and chinook
salmon (Burzawa-Gerard, Fontaine, 1965; Donaldson, Yamazaki, 1968; BurzawaGerard, 1971; Donaldson, 1973) on various species of fish (intact and
hypophysectomized fish) has shown that these preparations are widely effective, despite the fact that they were prepared from pituitaries of fish close
to genital maturity.
For example, administration of a salmon gonadotropin
stimulates all growth phases of genital cells (including spermatogenesis,
spermiation, vitellogenesis and ovulation).
These data corroborate the
existence in the pituitary gland of these fish of one gonadotropic hormone
with a multipurpose effect.
This is also suggested by the above data on the
possibility of stimulating the various gametogenic stages in intact specimens
by administering pituitary preparations of fish of the same stage of maturity.
The data that suggest that the pituitary gland of fish secretes two gonadotropic hormones are less convincing.
However, a final resolution of this
problem requires more research, particularly with respect to clarifying the
chemical nature of the gonadotropic hormones of Lower vertebrates.
In the more highly-organized vertebrates--the amphibians--hypophysectomy results in terminating vitellogensis and spermatogenesis. Proof that
the gametogenic and endocrinic tissues of the gonads in amphibians are controlled by the pituitary gland has also been obtained by administering
gonadotropic hormones.
Administration of purified mammalian FSH and LH to
separately hypophysectomized Rana temporaria (Lofts, 1961) and Rana pipiens
produces different effects.
LH fails to induce the development of either
spermatogonia or Sertoli cells, but it does affect the interstitial cells and
induces the spermiation reaction.
FSH, on the other hand, fails to induce
this reaction, nor does it affect the condition of the interstitial cells,
/41
15
but it does stimulate spermatogenesis and Sertoli cells (Dodd, Wiebe, 1968).
It is important to note that, in fish, FSH had no effect on the course of
spermatogenesis, whereas in amphibians the effect on spermatogenesis cannot
be doubted.
In this regard, the amphibians apparently are closer to the
Higher vertebrates. These data suggest that the pituitary of amphibians
secretes two gonadotropic hormones which affect different areas.
This
assumption is supported by histochemical data according to which two types
of gonadotropic cells have been detected in the posterior lobe of the pituitary of amphibians, and each of these cells has a different functional
activity, depending on the various experimental effects (van Oordt, 1968).
However, the gonadotropic hormones of amphibians have not yet been
sufficiently described biochemically, and thus, despite the extensive data
supporting the presence of two gonadotropic hormones in amphibians, this
problem has not yet been resolved.
Reptiles have the same cycles, which undoubtedly require the presence
of gonadotropins to initiate them, as fishes and amphibians. There is no
doubt that two gonadotropic hormones, distinct in chemical and biological
properties, have been detected in the pituitary gland of reptiles. One of
these hormones seems to function much like LH in mammals, and the other is
similar to FSH (Licht, Papkoff, 1974).
Sexual hormones
The sexual steroids play a very important role in the reproduction
process. The general function of these hormones is to control the secondary
sex characters.
The latter do not develop after a gonadectomy, and if they
already existed, the operation causes them to regress.
In castrates the
secondary sex characteristics can be stimulated by means of androgens and
estrogens prepared either synthetically or obtained from sex glands.
Sexual
16
steroids are formed under the influence of gonadotropic hormones from the
pituitary gland (see above). Androgens and estrogens stimulate the development and maturation of sexual cells.
The other effects of the sexual steroids (participation in regulating
ovulation, spermiation, spawning behaviour, and so on), which will be discussed below, may, in part, depend on other hormones such as gonadotropin,
pituitary preparation, prolactin and neurohypophyseal hormones.
Information on sexual hormones can be found in a number of papers
(Dodd, 1955, 1960; Hoar, 1955, 1965; Pickford, Atz, 1957; Ball, 1960;
Marschall, 1960; Forbes, 1961; van Oordt, 1963; Barr, 1965, 1968; Nandi, 1961;
Lofts, 1968; Hoar, 1969; Liley, 1969; Idler, 1972; Turdakov, 1972; Barannikova, 1973; Kirshenblat, 1973, and others).
A bibliography on steroid hor-
mones in fish is given in the index of Bern and Chieffi (Bern and Chieffi, 1968).
There is an extensive literature on the influence of sexual hormones
on sex differentiation and on the redefinition of sex in cold-blooded vertebrates. These problems deserve a special discussion, but that is not the
task of the present paper. The main points of these problems are discussed
in a number of papers (Burns, 1961; Forbes, 1961; Yamamoto, 1969; Reinboth,
1970).
In mammals, pregnenolone and progesterone, which are formed from
cholesterol, are the main steroids that initiate the synthesis of androgen
and estrogen.
In the Lower vertebrates pregnenolone is also central to the
biosynthesis of steroid hormones.
Progesterone, which is initiated in several
intermediate stages by cholesterol, forms the underlying base for androgens
(with testosterone as its main link) (Hoar, 1965, 1969; Yamamoto, 1969).
These metabolic chains are very old phylogenetically. Progesterone, estriol
1713 and other estrogens have been identified in the ovaries of a number of
17
invertebrates and they also occur universally in vertebrates, although they
have a certain specificity in cold-blooded vertebrates (Hoar, 1965; Barr,
1968).
Estrogen-like compounds have also been detected in plants (Bickoff,
1963).
The functional structures that produce the steroids are identified
and studied by means of various physiological, biochemical and histochemical
methods.
The data obtained from enzyme studies are particularly important--
enzymes that are linked to the biosynthesis of steroids (primarily 3^hydroxysteroid dehydrogenase 3S HSD) in the sex glands (Hoar, 1969). The
testis of cold-blooded vertebrates contain interstitial cells (comparable
to Leydig cells of the Higher vertebrates) that contain lipoid inclusions
and cholesterol.
Moreover, in a number of fish the walls of the seminal ducts
contain boundary cells with similar inclusions in the cytoplasm; these cells
also produce steroids and, in a physiological sense, are homologs of the interstitial cells (Lofts, 1968).
The testis of most Lower vertebrates contain
Sertoli cells which, according to a number of investigators, also participate
in the production of steroids. The steroid-producing cells in the testis
of Lower vertebrates function according to a precise seasonal cycle linked
to the reproduction process and to the development of secondary sexual characters.
No such precise data are available with respect to the structureS
that generate the ovarian hormones.
On the basis of histochemical methods
it is assumed that estrogens are generated by granulose cells and by the
follicular theca (Bara, 1965; Lamber, 1966; Barr, 1968). These data suggest
that the follicular membranes of the developing oocytes contain enzymes which
are involved in the production of steroids. The role of granulose cells
18
(follicular epithelium) is emphasized in most cases with respect to these
processes in fish (Lamber et al., 1972).
By using a number of rigid standards it has been demonstrated that
steroids occur in ovarian and testicular tissues and in peripheric blood
of the major groups of the Lower vertebrates. Studies have also been made
of the enzymes that figure in the synthesis of steroids, and a number of
steroids have been synthesized in vitro from the tissues referred to above
(Ozon, Breuer, 1963;1dler, Truscott, 1963; 1966; Breuer, Ozon, 1965; Barr,
1968; Lofts, 1968, and others).
These data suggest that the sexual glands of Lower vertebrates synthesize sexual hormones which take part in the endocrine system that controls
reproduction (Barr, 1968; Fontaine, 1969; Lofts, 1968; Hoar, 1969).
Male Sexual Hormones
Cyclostomes. Interstitial and boundary cells containing lipoids
and cholesterol have been detected in the testis of the oldest Agnatha
vertebrates.
The clear-cut seasonal cycle of these cells is linked to the
reproduction period (Marshall, 1960; Larsen, 1965).
Growth of secondary
sexual characters was obtained experimentally (swollen and hyperemic cloacae
in lamprey) under the influence of testosterone.
Gonadectomy in lampreys
prevents the growth of secondary sexual characters.
This suggests that
their androgens are being generated by the endocrine tissue of the gonads,
for in these animals androgen secretion is already regulated by the pituitary,
since the latter's removal results in the extinction of secondary sexual
characters (Dodd, 1960; Larsen, 1973).
19
Elasmobranchs. In this group of fish the androgens are being
secreted in the testis.
Administration of androgens resulted in increased
growth of the claspers in the shark Mustelus canis (Pickford, Atz, 1957).
The presence of steroid-producing tissue in the testis of these fish has
And enzymes linked to
been established histochemically (Chieffi, 1962).
the generation of steroids have been found in Sertoli cells (Simpson, Wardle,
1967).
The steroids (testosterone, estradiol l7
,
progesterone) were isolated
chemically (Chieffi, 1962, 1967; Hisaw, 1963). Testosterone was determined
in the blood of a male Raja radiata (Idler, Truscott, 1966). Apparently
testosterone is the main androgen in the elasmobranchs.
Teleostomi. The effects of androgens are quite varied in the teleosts.
Removal of the testis results in lack of development of special coloration
linked to sexual dimorphism; loss of breeding color.
The special formations
(pearly rash, gonopodia in a number of species) that are associated with the
reproductive period in fish fail to appear if the testis is removed. These
and other secondary sexual characters develop in castrates and intact young
fish when they are administered doses of testis tissue or androgens (Dodd,
1955; Pickford, Atz, 1957; Forbes, 1961; Hoar, 1965, 1969).
Castration disturbs the spawning behaviour in many fishes.
For
example, the castrated stickleback male doesn't build a nest, and progeny
care also depends on androgens (Hoar, 1962).
The behaviour of male stickle-
backs is quite varied during the spawning period. It turned out that the
androgen level of dominant males was 5--7 times higher than that of the less
dominant ones.
terone.
And only the testis of the dominant males contained testos-
(Gottfried, van Mullem, 1967). Administration of testosterone to
castrated gourami males resulted in a regeneration of their secondary sexual
characters and in characteristic spawning behaviour (Jones, Liley, 1970).
20
Common paradise fish will start the initial nest-building phase under the
influence of androgens but they will complete the process only if prolactin
is administered (Machemer, Fidler, 1965).
Thus androgens in combination with other hormones and neurohormones
play an important role in the adaptive spawning behaviour peculiar to the
species (Baggerman, 1968; Liley, 1969).
But not all data obtained in this
connection were identical: Tilapi macrocephala, in particular, continued
with nest building even after the gonads had been removed (Hoar, 1965).
Androgens are very important in stimulating spermatogenesis and
spermiation; some of this information was already discussed above. Prolonged
administration of testosterone-propionate to hypophysectomized male Fundulus
heteroclitus stimulated the interstitial cells and increased spermatogenesis
(Lofts et al., 1966; Pickford et al., 1972). Testosterone-propionate also
stimulated spermatogenesis in the hypophysectomized small catfish Heteropneustes fossilis (Sundararaj, Nayyar, 1967).
Detailed studies have been made of the steroid-producing cells found
in the testis of many teleost species and they have disclosed the enzymes
that participate in the synthesis of steroids (Bara, 1966, 1969; Yaron, 1966;
Wiebe, 1969; van den Hurk, 1973). Still, the 3S HSD-activity occurs in
various structures of various fish species. In most of the fish in question
the steroids are synthesized in the interstitial cells (Marshall, 1960; Lofts,
Marshall, 1957; Ahsan, 1966; Lofts, 1968; van den Hurk, 1973). These cells
are considered to be the main source of the sexual steroids in the testis of
teleost fishes.
Moreover, the enzymes that participate in the biosynthesis
of steroids were detected in the boundary cells of ducts (Tilapia mossambica,
Yaron, 1966; S. salar, 0'I-Ialloran, Idler, 1970).
In some species Sertoli
cells contained 3G HSD (Bara, 1969; Wiebe, 1969), but in other species these
/44
21
cells revealed no steroid synthesis (van den Hurk, 1973). In many teleost
species sexual steroids were determined in the tissues and in the blood,
and the quantity of these steroids increases in proportion to gonadmaturation
(Hoar, 1969).
In amphibians the Sertoli cells were found to have steroidal
properties (van Oordt, 1970).
As in amphibians, 3 HSD-activity has also
been detected in the Sertoli cells of reptiles (Lofts, 1972).
Female Sexual Hormones
Estradiol 17S was identified in the ovaries of various taxonomic
groups of fish and fish-like organisms. This steroid has also been detected
in a number of invertebrates. Extensive distribution of estradiol 170 in
various animlas underlines the phylogenetic origin of this hormone. It may
be assumed that, physiologically, estradiol 17S is the most important estrogen.
Many other, less distributed estrogens also originate from estradiol 175.
Progesterone has been identified in the tissues of all vertebrates and many
invertebrates.
Progesterone is an important link in the synthesis of steroids
and it is an important hormone with a distinct physiological effect of its
own (Hoar, 1965, 1969).
Cyclostomes.
Ovariectomy in Petromyzon fluviatilis prevents the
growth of secondary sexual characters in this species. Hence the ovaries of
the oldest vertebrates secrete sexual hormones with a function close to that
of the-more complex animals (Dodd, 1960; Hoar, 1965).
-Admi.zqlstration of estradiol to intact female lampreys increases the
calcium content in the blood. Owing to the greater synthesis of calcium-rich
phosphoproteins in the liver, which are incorporated into the developing
oocytes, the oocytes increase considerably in weight (Pickering, 1973).
22
Progesterone, estradiol 178 and estrone, which are important hormones
in the biogenesis of genes (Hoar, 1965), have been found in the ovaries of
Petromyzon. The question regarding the localization of steroid secretion
in the ovaries of cyclostomes has not yet been completely resolved. Based
on data pertaining to the ultrastructure, estrogen is assumed to be secreted
The greatest 38 HSD activity
in the follicular theca (Busson-Mabillot, 1967).
was detected in the follicular cells (Lampetra planeri, Hardisty, Barnes, 1968).
Elasmobranchs. Estrogens and progesterone have been identified in
the ovaries of this group of fish (Simpson et al., 1963; Chieffi, 1962--report on steroids identified in elasmobranchs).
It was shown that the amount
of estrogens increases with oocyte maturation, based on Scyliorhius caniculus.
A number of steroids have been detected in the blood plasma of Torpedo mormorata at various stages of the sexual cycle (Lupo et al., 1967). Administration of estradiol to female Scyliorhinus caniculus at various stages of
maturation induced oviduct development only in specimens close to maturity
(Dodd, 1960). Administration of estrogens induces development of the female
sexual tract in oviparous species (Chieffi, 1967).
The location of estrogen secretion in elasmobranchs is not clear. It
has been demonstrated that the ovaries of
Raja
crinacea and Squalus acanthias
LO
have the abilityeransform pregnenolone into progesterone (Chieffi, 1966).
The yellow body that forms in the ovaries of a number of sPecies of selachian
fishes is not regulated by the pituitary, as in the Higher vertebrates, and
hypophysectomy does not interrupt pregnancy in Mustelus canis (Hoar, 1965).
However, the work of other investigators has shown also that in Torpedo
marmorata the
corpus
luteum acquires an epithelial character and that it is
a gland connected with the production of steroids.
This is based on histo-
chemical studies and, in particular, on the fact that progesterone has been
23
identified in the yellow body (Chieffi, 1962; di Prisco et al., 1965).
-Teleostomi. Ovariectomy results in involution of secondary sexual
chhracters in telost females. Gonadectomy in Xiphophorus helleri and Betta
splendeus obliterated all sexual activity (Dodd, 1955,1960); in Tilapia it
resulted in a change in breeding color, but this effect was not observed in
Amia calva and Gasterosteus aculeatus (Dodd, 1955, 1960; Ball, 1960). The
growth of the ovipositor is a convenient measure of the estrogen activity
in the bitterling Rhodeus amarus and R. ocellatus (Bretschneider, Duyvene de
Wit, 1947; Shirai, 1962). Ovariectomy in Lebistes reticulatus during pregnancy reduces the epithelium of the oviduct, and administration of estradiolbenzoate prevented this involution (Chambolle, 1965). Ovariectomy in gourami
females resulted in extinction of spawning behaviour. When these females were
placed together with males which were in spawning
condition, the ovariectomized
females attracted the males less than the intact females. It is thought that
the ovary, directly or indirectly, controls the secretion of the chemical
substances that attract the male (Liley, 1969).
Although this aspect of the
sexual steroids is very interesting, it has been investigated relatively little
(Fontaine, 1972).
Estrogens are very important for the development and matura-
tion of oocytes. In fish and amphibians estrogens induce the synthesis of
calcium-rich phosphoproteins in the liver which participate in forming the
yolk of the growing oocytes.
Sturgeons, teleosts and amphibians have provided interesting data on
the progesterone mechanism.
The effect of progesterone on the in vitro ovary
of sturgeons is that meiosis can be induced in the eggs that have been released
from follicular epithelium. Hence oocyte maturation must occur under the
influence of progesterone.
The gonadotropic hormones of the pituitary in
these fish, as in amphibians, affect the cells of the follicular epithelium,
24
stimulating these cells to secrete a progesterone-like substance that subsequently causes the ovicells to mature (Detlaf and others, 1968; Goncharov,
1969; Detlaf, 1970).
The significance of steroids, which are being synthesized in the
ovary under the influence of gonadotropins, for oocyte maturation has also
been demonstrated in different species of teleost fishes (Jalabert, Breton,
1973).
Small doses of progesterone (0.25 mg) induce ovulation in Carassius
auratus (Yamazaki, 1965), and a similar effect is obtained in loaches
(Kirshenblat, 1952).
Estradiol ln, estrone, estriol and progesterone have been indentified in the ovaries and in the blood of teleosts. The content of these
substances varies according to season, and estriol appears in significant
quantities only during the spawning period (Ball, 1960; Barr, 1968). In
salmon the estrogen content fluctuated seasonally in both males and females.
The highest estrogen content occurs during the spawning period (Cedard et al.,
1961).
The 3(3 HSD activity in the ovary of Brachydanio rerio during the
reproductive cycle is also of interest.
The enzyme occurs in the cells of
the follicular epithelium during vitellogenesis and also in the corpus luteum
after ovulation. The production of steroids is at its height after the spawn.
These changes coincide at this period with increased activity of the gonadotropic cells of the pituitary (Lambert et al., 1972). These data are interesting from the point of view of the reciprocal relationship between the
hypothalamus-pituitary complex and the gonads in fish.
In amphibians, as in other vertebrates, the steroids are.synthesized
in the ovary. Extracts obtained from ovaries contain progesterone, estrone,
estradiol 1713 and estriol (Barr, 1968).
25
In reptiles the ovary contains corpus luteum which survives during
the entire pregnancy period and is reminiscent of the corpus luteum of
mammals.
It is assumed that this body has an endocrine function; it contains
progesterone (Forbes, 1961).
Thus the effect of the ovarian hormones in reptiles differs somewhat
from that of the Lower vertebrates and is reminiscent of that found in the
more highly organized animals.
Conclusion
These data illustrate the complex and peculiar interaction of the
gonadotropic and sexual hormones in the regulation of the reproductive system
in various classes of cold-blooded vertebrates.
In the oldest Agnatha vertebrates there is no precise pituitary
control of the development of sexual cells--something that occurs in all
Higher vertebrates, beginning with fish.
In the cyclostomes the gonadotropic
hormone of the pituitary undoubtedly regulates the steroidal tissue of the
gonads.
The effect of the sexual steroids in the cyclostomes is varied and
similar to that found in fishes.
The early hypophyseal control of the secretion of sexual hormones
in the phylogeny apparently is based on the need for a more reliable effect
of these hormones.
In all groups of.fish (beginning with.cartilaginous fish), as in the
Higher vertebrates, the gonadotropic hormones of the pituitary are needed
for gametogenesis and for maturation of the sexual cells, with the regulating
hormonal effect being most prominent at certain periods. The most important
function of the gonadotropins is to stimulate the endocrine tissue of the
gonads to secrete sexual steroids.
The effect of the gonadotropins on the
sexual cells of certain growth phases consists in that this hormone induces
26
the secretion of steroids which bring about the appropriate effect.
A consideration of the effects of the gonadotropic hormones on the
reproductive system requires data on the number of gonadotropic hormones
and on the different effects they have in various cold-blooded vertebrates,
compared to the effects of LH and FSH in the Higher vertebrates.
As to the cyclostomes, we have not yet obtained precise data on the
number of gonadotropic hormones in the pituitary. Endocrinologists are also
occupied with this problem.
The pituitary of many fish species (but not of
all) has been described as having two types of gonadotropic cells; however,
the sexual glands of hypophysectomized and those of intact fish can be
stimulated at various stages of the cycle (vitellogenesis, spermatogenesis,
meiosis, ovulation, spermiation) with the same pituitary preparation obtained
from fish specimens with close to mature sex glands. This effect was obtained
by administering either acetonized or fresh pituitary preparations as well
as by using a preparation of purified fish gonadotropin.
These data suggest that the fish pituitary secretes one gonadotropic
hormone with a multipurpose effect. This is confirmed to a certain degree
by research on the effect of LH in fish, namely, that the use of LH in
hypophysectomized specimens stimulates both spermatogenesis and the steroidal
tissue (FSH does not produce a similar effect). Thus the LH of mammals has
wide-ranging effects in fish and in the number of cases its effect is similar to that of FSH (in Higher vertebrates). However, some data suggest that
the fish pituitary contains either two gonadotropic hormones, or one hormone
with two fractions.
This problem requires further research.
Amphibians are more likely to have two gonadotropic hormones. In
these animals LH does not induce spermatogenesis but it does stimulate the
interstitial cells (as in the Higher vertebrates). FSH is effective in
amphibians (in contrast to fish) and induces spermatogenesis. These interactions
/47
27
between the gonadotropic and sexual hormones have a greater similarity to
those in the Higher vertebrates than to those in fish. In reptiles the
existence of two gonadotropic hormones has been demonstrated and their
effect is similar to that of LH and FSH in the Higher vertebrates.
The sexual steroids play a very important role in the development
and maturation of the sexual cells and in the reproduction process.
steroids have a long phylogenetic history.
Many
This applies particularly to
estradiol ra and progesterone, which occur in all vertebrates and in some
invertebrates.
The biosynthesis of the different steroids is also very
similar.
Progesterone is an important link in the biosynthesis of steroids.
However, this hormone also has a distinct and significant physiological
effect.
It is progesterone (or a progesterone-like substance), synthesized
under the influence of a gonadotropin, that induces oocyte maturation (in
teleosts, sturgeons and amphibians).
Testosterone is the universal hormone of males. It turned out that
spermatogenesis in fish is induced not only by gonadotropins but by testosterone as well. The effect of sexual hormones on the development of secondary sexual characters has been investigated in greatest detail in various
classes of vertebrates.
The occurrence of species-specific, genetically
determined spawning behaviour is largely influenced by sexual hormones acting
synergically with a series of other hormones and neurohormones.
There has also been clarification with regard to another steroidal
effect, involved in inducing spawning behaviour and in maintaining a particular hierarchy with adaptive significance for the fish population. The sexual
steroids in fish induce the secretion of substances (apparently ectohormones
[sic] or pheromones) that attract specimens of the same species but from the
28
opposite sex.
It is this (along with other factors) that synchronizes the
reproduction process in the males and females of a given species, which is
an important adaptive mechanism that facilitates reproductive success and
survival of the species.
29
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.
S a p,a H H H x o B a Il A. KoxueHrpau}!st roHaaorponHoro roptona B rnn0^H3e CaMüOB H
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•
•
Gonadotropic and sex hormones and their 'role in the regulation of the reproductive .
in poikilo thermic vertebrates
sytem
I. A. Barannikova
4
SUMMARY
The role of gonadotropic and ,sex hormones in the regulation of gonads function
and reproduction in fishes is considered. The data in this relation on the Agnatha
and higher classes of cold—blooded vertebrates are shortly considered. The development of the regulation of the reproductive system in different groups of poikilothermic
vertebrates is discussed on basis of these data.