The malaoologioalmalacological societysooiety of Japan
貝 雑 VENUS （Jap． Jour． Malac ．） −． Vo1 ，48， No ．2 （1989 ） ： 99 139
総 説 Review
陸 生 貝 類 に お け る 生 殖 の ホ ル モ ソ 支 配
武 旧 直 邦 ． 〔東邦 大学 理 学 部 生 物学 教室 ）
Hormonal Co1ユtrol of Reproduction in Land Snails
Naokuni TAKEDA
〔Department of Biology ， Faculty Qf Science ， Toho University ，
Funabashi ， Chiba 274， Japan）
CONTENTS
I． Introduction ．．．．．．．．，．，，．．▼，，．．．、。，， ．100
1L Neuroendocrine Systems ．．．．．，．．．，．．． ．101
A ． Neurosecretory Systems ．．，，，，．．．，， ，101 B ． Endocrine Organs 、．．．．．．．．．．．．．．．． ．102
a ▼ Dorsal bodies ．．．．▼，．．．，．．．．．．．．．， ．103
b『 Optic glands ．．．．、．．．，，，．．，．，．．．．． 。104
c ． Hermaphrodite glands ．．，．．．．．．．．． ．106
11L Hormonal Control of Reproduction ．．．．，．．．．，．．．．．．．．．．，．，．．．．．．．106
A ． General Morphology of the Reproductive System ．．．．，．．．，．．．．．．106
B ， Maturation of the Reproductive System ．．．，．．．．，．，，．．．，，．．，．．107
a ． Ga 皿 etogenesis ．．．．．，マ．．．．．．，，，．，．．『，呷。．．．．．．，，，，．．，．．．．．『．107
b ． Accessory sex organs ．，，．．．．，，，．．．，．．、．，．．，．，，．．，。．．，，，．，，111 1， Head −warts ．．．．，．．辱，．『．，凸．，レ，．．，，．．，呷，．，．．．の．．．，．『疊．．，．112
2． Albumen glands ．．，，．．，，，．．．．，．，，．，．，．，．．．，．．．．．．．，．．．．113
3． Reproductive tracts ．．．．．．．，，．．．．，．．，．，．． ．，．，．．，．．．，．．．．113
C ． Mating Behavior ．．．．．．．．．．．，．．．．＿ ．，．．，，，．．．，．，．．．．，．．．，，．115 D ． Oviposition ．．．呷．．．．．，．，．呷．．．．．『．，．．．，．．，，．，，．，．．．．．．．．．．．．．118
1V ， Dynamics of 且 ormonal Control of Reproduetion ．．．，．．．．，．．，．．．，，119
V ． Comparative Endoerinology of Reproduction in Molluscs ．．．．．．．．．，121 References ，．，．．．．，．．凾．．．呷．幽．，．．．．．．，．鹽鹽．．．．，．．，．層．．．，．．．．，．．，．．．，，128
一 NII-ElectronicN 工 工 Eleotronio Library Service The malacologicalsocietymalacological society of Japan
100 VENUS: VoL48, No. 2 (1989)
Neuroseeretory cells oeeur in the eentral I. Introduction nervous system in all molluscan speeies.
Mollusean species show a great diver- In the less organized groups such as
sity. They are adapted to a variety of Bivalvia or Prosobranchia, epithelial
different environmental conditiens. The endocrine glands are entirely lacking,
greatest ranges ef environmental con- as feund in the lower invertebrates ex-
ditions that must be survived by organ- cept the Arthropoda. Therefore, neuro-
isms are those found on land. Terrestrial seeretions are the only chemical eo-
forms are found only in prosobranch ordinators in these speeies. With the 'ehese, and pulmonate gastropods, Among inerease in body eornplexity, endoerine
pulmonates are far more suceessful on glands, such as the dorsal bodies and
land than prosobranchs. In particular, the optie glands, are developed in highly
stylommatophoran species are exelusively erganized speeies such as pulmonate
terrestrial and are considered to have gastrepods and cephalopods. Thus, the
adapted to the land habitat directly neuroseeretory system and the endoerine
from the sea, as they have no fresh- glands so eoordinate reproduction that
water aneestors (Joosse, 1979). Slugs mollusean speeies ean propagate them-
are eonsidered to have evolved from selves functionally.
snails in whiah the shell has degenerated Our knowledge of the hormonal eon-
as they have a eommon developmental trol of reproduction en molluses has
stage in whieh a cap-shaped shell is inereased (e.g. Boer and Joosse, 1975;
formed. Takeda, 1981: Joesse and Geraerts, 1983;
With the adaptation to life on land, a Geraerts et al,, 1988; Joosse, 1988).
number of physiological ehanges have However, the studies are mainly re-
occurred. These are lung respiration, strieted to those on Lymnaea highly developed senee organs, a eon- water basommatophoran pulmonates> eentrated nervous system and complex (e.g. Geraerts and Joosse, 1975; Joosse, reproduction, As to reproduetion, all l975; Joosse et al. 1985). ApZysia (ma- pulmonates are hermaphrodites, and the rine opisthobraneh gastropods> (e.g. male and female gametes differentiate Strumwasser, 1984; Arch and Gainer, within a single gonad, called the herma- 1985; Geraerts et al., 1988), and Oetopzts phrodite gland This is different from (cephalopods) (e.g. Wells and Wells, other hermaphrodite animals where 1977; Wells, 1978) and there are cem- there are two different gonads or one paratively few in studies on stylommato- sex succeeds another (Reinboth, 1975). pheran pulmonates (e.g. Gomot et al., All stylommatophoran speeies are true 1980; Takeda, 1982a; Takeda, 1985a; simultaneous hermaphrodites, although Takeda et al., 1986; Gomot and Delay, they may show functional protandry 1987), Therefore, in the present Teview, (Tompa, 1984). In addition, self-fertili- the hormonal control of reproduction of zation and parthenogenesis are also land snails and slugs is diseussed from found in some speeies (Ikeda, 1937; the view point of eomparative endocri- Emura, 1981). nology. The rnain rnaterials are the The reproductive process is generally snails, Euhadra peliotnphala･ and Acha- known to be under the hormonal control, tina fulica, and the slugs, Limax margi- NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda: Hormonal Control of Reproduction in Land Snails 101 natus and DeToce7'as reticulatnm which Ktihlmann, 1964; Gabe, 1966; KUhlmann, are commonly available in Japan. 1966; van Mol, 1967; Boer and Joosse, 1975; Kai-Kai and Kerkut, 1979; van II. Neuroendocrine System Minnen and Sokolove, 1981; Roubos, The endocrine systems serve as bases 1984; Ridgway, 1987). These neuro- foT the control of reproduction, Gen- seretory eells are often unipelar and erally, neurosecretory cells in mollusean show a great variability in size. The speeies are grouped within ganglia and nuclei are lobed and contain large num- are assoeiated with the regions of the ber of nucleoli. Sometimes, they con- perineurium of nerve adjacent to blood tain many inclusions sueh as glycogen, vessels, forming neurohemal areas. His- lipid droplets, lipofuscin and carotenoids, tolegieal changes in these cells are cor- They are located at the periphery of related with a variety of reproduetive each ganglion. A large group of neuro- phenomena. Among invertebrates, the secretory cells is situated in the central species of molluscs and arthropods are and dorsal parts of the mesocerebrum the most highly organized in body struc- apposed to the perineurium elosed to tures and have some endoerine glands. the origin of the eerebral commissure. Many neurohormones released from The axons of these cells form a bundle neurosecretory cells aet direetly on spe- whieh passes into the eerebral neuropile, cifie target tissues, whereas others con- where it branches into a larger and a trol activities of endoerine glands in smaller one. The entire peripheral re- secreting hormones, Thus, in eombina- gion of the nervous systern is used for tion with the neuroseeretory system, the hormone release in most molluscan spe- endocrine glands form an endocrine eies, this is termed peripheral neuro- system whieh in arrangement and scope secretlon. resemble those of vertebrates. Surpris- In Ezthadra peliompha･la and Acha･tina ingly, molluscan hermaphrodite glands fulica, the brain is eovered with the are known to secrete sex steroid thick and hard connective tissue dif- hormones whieh play a similar role in ferent frorn that ot Limax marg･inatus. principle to those of vertebrates. Neuroseeretory eells in the eerebral ganglia were found in the medial, lateral A. Neurosecretory Systems and posterior parts forming a mass. In The central nervous system in mol- these species, medial neurosecretory cells lusean species consists of a number of ehange aetivity in relation to reproduc- ganglia eonnected by commissures and tion: during the breeding season, they conneetives. These are the paired eere- beeome aetive in the produetion and bral (brain), pleural, parietal, pedal release of seeretory materials whereas ganglia and a single visceral ganglion, outside the breeding season their ac- Sinee the first dernonstration of neuro- tivity beeome low. Fine strueture of secretory eells in the Mollusca by the aetive medial neurosecretory eells in SchaTrer (1935) in Pleurobranehea (ma- Euhadrq petiomphala is shown in Fig. rine opisthobranch gastropods), neuro- 1. Many well developed Golgi apparatus, seeretory cells have been deteeted in all which indicate the produetion of seere- groups of molluscs (e,g. Nolte and tory granules, are seen. Neuroseeretory NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 102 VENUS: Vol.48, No. 2 (1989) Fig. 1. Fine structure of the medial neurosecretory of the brain in the snail, -v -? E"hadra PeliomPhala in the breeding season,cells tkMWIo y t.p ( ac ss et6 ma op ･pa me 5> mu pm te oMmaMk,E, materials aTe known to be released fTom has been demonstrated in the nerve cells the cell body or the axonal endings into (Schot et al,, 1981; Joosse and Geraerts, the hemolymph via the neurohemal areas. 1983; Kobayashi et al., 1984; van Noor- In Euhadra peliompha･la and Limax den, 1984; Marchand and Gomot, 1986). marginatt{s, the majority of the neuro- In the present speeies, many neuro- secretory axons run to the neurohemal peptides have been detected in the area, the peripheral regions of the central nervous system (Takayanagi ganglia and others innervate directly the and Takeda, 1984; Takeda et al,, 1985; target organs sueh as the optic gland Fukutomi et al., 1986; Takeda et al., and the dorsal bodies as described later. 1986; Takeda, 1987a; Takayanagi and In the snail, Helix ponati.a, neuroseere- Takeda, 1987; Mizuno and Takeda, tory materials are released into the 1988a,b; Takayanagi and Takeda, walls of the cerebral arteries (Nolte and 1988a,b). These are FMRFamide, arg- Ktihlmann, 1964; van Mol, 1967>, vasopressin, oxytoein, met-enkephalin, The classie picture of the neuresecre- pancreatic polypeptide, insulin and so tory system has been ehanged by the on. However, at the present time, their advance of immunohistochemistry. The participation in reproduetion is unclear. presenee of substanees reaeting with antibodies against the biologically aetive B. Endocrine Organs peptides, including vertebrate hormones In phylogenetieally lower species, NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda: Hormonal Control of Reproduction in Land Snails 103 endocrine glands are absent, In sueh S?tccinea putris (Lever, 1958; Cook, speeies, endocrine eentrols are per- 1966), Strophocheilus oblengus (KUhl- formed simply by neurohormones se- mann, 1966>, Acavis phoenix (Boer and creted from neurosecretory cells. Only Joosse, 1975) and Achatina fulica in gastropods and cephalopods, endo- (Ohtake and Takeda, 1987). Thus, de- crine organs play an important role in velopmental states of the dorsal bodies the control of reproduetion, usually differ from species to species. In the under the direetion of neurohormones. present species, the dorsal bodies are a, Porsal bodies well developed in Deroceras reticulatum The dorsal bodies are the endoerine and Achatina fulica. On the eontrary, organ attaehed to the perineurium of their state is unclear in Euhadra peliom- the brain. Lever (1957, 1958) was phta and bimam marginatus. Fine struc- the first to suggest an endoerine Tole ture of the dorsal body eells in Achatina for the dorsal bodies in freshwater .futica is shown in Fig. 2. In this species, snails. The oecurrenee of the dorsal two types ef granules are found: ene is bodies in stylommatophoran species was large, with a thick electron dense eortex, first reported by Nolte (1965), In con- and the ether is small and arised from trast to the situation in basommato- the Golgi apparatus. The number of pheran species, the dorsal body cells in large granules inereased in the breeding stylommatophoran speeies are dispersed seasen, suggesting their participation in in groups in the eenneetive tissue sheath reproduetion. In these eells, mitoehon- around the brain (Nolte, 1965; Cook, dria "'ith well developed cristae are 'the 1966; Ktihlmann, 1966; van Mol, 1967). characteristically numerous, In Compact dorsal bodies have been ob- eonnective tissue of the medial part of served in only a few snail such as the brain, a bundle of commissural Fig. 2.Fine structure of a dorsal body cell in the snail, Achatina frtlica, Scale, O. 5 "m. -v 7 7 v rb -d i v- tisO6??ENfifgnvrzoikanssjli. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 104 VENUS: VoL 48,No, 2 (1989) nerves, whieh also contain neurosecre- et al., 1986) and to be immunoreaetive tory granules, is found, Furthermore, for met-enkephalin in Helix aspersa synapse-like structures are also found (Marehand and Dubois, 1986). In addi- between the axons of commissural nerves tion, the gland eells have eilia, assu- and the dorsal body cells. Unknown in ciatred with a basal body and a proximal basommatophoran species these innerva- eentriole, The dorsal body cells seem to tions have been found in other snails, originate from ciliated epithelial eells such as Theba pisavea (Nolte, 1978) and with a sensory nature. The cilia are Helix aspersa (Vincent et al., 1984>. also found in the dorsal bodies of Hetix Therefore, the dorsal body eells of land aspersa (Griffond and Vineent, 1985) stylommatophoran pulmonates are sug- and the juxtaganglionar organ, a homo- gested to be under the control of neuro- logous organ to the dorsal body, of the seeretion originating frorn the brain. trochid Gibbula umbilicalis (proso- In Helix aspersa･, the dorsal bodies have branchs) been shown to be under the inhibitory brates, ciliated endoerine eells have control of the brain (Wijdenes et al., generally been found in the hypophysis, 1987). The increases in eell size and the medulla and the pancreatic islets. protein synthesizing aetivity have been b. Optic glands "optie found at the egg-laying period (Griffond The name gland" was originally and Vineent, 1985). The ehemieal applied to the endoerine gland of eepha- nature of the dorsal body hormone lopod$ (Wells and Wells, 1959). The has been suggested to be steroid, same name is also used for the tentaeular ecdysone, in the snail, Helix pomatia hormone seereting glands in stylommato- (Krusch et al., 1979; Nolte, 1983; Nolte phoran pulmonates (Takeda, 1982b). Fig･3. Fine structure of the optic gland cell including innervation (insert figure) in the snail, Euhadra PeliomPhata, -vf s .z l7I vcs"o-6unwamteoanmauristh, .".E:a)npal･:, km tets"opaffhfi6r7St. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda: Hormonal Control of Reproduction in Land Snails 105 Pelluet and Lane (1961) were the first postulated not only to end in the epithe- to suggest an endoerine tunction for lial surface but also to end in surround- the optic tentacles of stylommatopheran ing blood vessels or in ether tissues speeies, C'ontradietory effects of ex- (Pelluet and Lane, 1961), Furthermore, tirpatien of the optic tentaeles on repro- the optic gland in the optie tentacles ductive funetion have been reported has been demonstrated to have a stimu- (Bride and Griffond, 1981). In many lating effect on spermatogenesis in vitro species of land stylommatophoran spe- and no hormonal aetion was found in eies, however, the existenee of a sexually- any region of the tentaeles other than controlling tentacular hormone has been the optic gland (Takeda, 1982b; Taka- well demonstrated (Pelluet, 1964; Berry yanagi and Takeda, 1985). In relation and Chan, t968; Gottfried and Dorf- to this, it is interesting to note that man, 1970a,b; Nagabhushanam and exocrine substances from the atrial Kulkarni, 1971; Wattez, 1973, 1975; gland of the reproductive tract in Sukumaran and Suriramulu, 1977; Ta- Aplysia californica, eontains peptides keda, 1977; Takayanagi and Takeda, which induce egg-laying when injeeted 1985; Wattez, 1978). As a result, four into mature animals (Arch et al., 1978, cell types have shown to be located near 1980; Sehlesinger et aL, 1981). The the tentaeular ganglion (Lane, 1964; optie gland cell has a long proeess and Rh6nlieh and Bierbauer, 1966; Bierbauer the eytoplasm is fi11ed with large gran- and T6rak, 1968; Rogers, 1969). These ules with a thin coTtex in Limax margi- are the optie gland cells (a collar cell natus, and with a thiek peripheral layer eluster), two types of lateral cells and in Euhadra peliemphala (Fig. 3) (Ta- the gland cells of the dermatomuscular keda et al., 1987). These gland cells layer. Three of these gland cells are are eharaeterized by a well-developed regarded as being subepidermal gland granular endoplasmic reticulum. Gran- cells while the optic gland cells have ule are formed in the Golgi apparatus, intimate connections with the tentaeular Although no neurosecretory cell bodies ganglion (Bierbauer and T6rdk, 1968; oecur in the tentacular ganglion, neuro- Takeda et al., 1987). In addition, the secretory axons penetrate into the optie eptic gland cell processes have been gland eells (Fig. 3: inserted figuTe). Table 1. Seasonal variations of sex steroid hormones in the hermaphrodite '(i gland in the snail, E"hadra PeliomPhala. i z y' yi ?if ontMiwa -e tialS 2t4tt-f p ･t r rti ,v ?olienesTt t tttt Steroid hermones testosterone progesterone estr5dioi (pg,,'mg tissue) t ttttt tt ttt t ttt Jan.Mar,MayJul.Sep,Nov, O.2 ± O.1 O.4 ± O.1 1.3 ± O.2 1120. 0± 144. 5 3224, 9± 433, 2 30. 0± 8, 6 3115. 1± 513. 2 7110. 6± 934. 4 180.2 ± 8.5 Months 4051. 3± 481. 0 482. 6± 687. 3 243. 5 ± 54, 3 72.1 ± 5.2 126.0 ± 13.4 28, 2 ± 8. 4 LO ± O,7 2, 1± O.8 4, 3± LO CMean ± S. D. : n=7) NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 106 VENUS: Vol. 48, No. 2 (1989) Cobalt fi11ing revealed that the axons of been shown to reduce remarkably, fol- rnedial neuroseeretory eells project to lowing castration. The amount of each the tentacular ganglion, near' the optic steroid hormone increased gradually at gland (Takeda et al., 1987), Therefore, the time of new gamete differentiation the optie gland is considered to be con- (Takeda and Sugiyama, 1984>. In these trolled by a neurohoTmone originating species, the flat cells, with a large from neurosecretory cells of the brain. nucleus and well developed smooth c. Hermaphredite glands endoplasmie retieulum, in the peripheral It is clear that in some stylommato- region of the aeinus may be concerned 'is phoran pulrnonates the gonad a souree with the synthesis of sex steroid her- of hormones. This is suggested by mones. In some species, there are sug- Abeloos (1943) and Laviolette <1954). gestions to synthesize steroid hormones They found from eastration experiments by the Sertoli cells (Serra and Koshman, in some slugs that the hermaphrodite 1967; Parivar, 1980). Many sex steroid the gland has an endoerine function on the hormones have been identified in et growth ef the accessory sex organs, In g'onads of Helix aspersa (le Guellee AgriolimQx reticulatus ticulatum), the proeess of maturation physiologieal stage and their physiologi- of the aeeessory sex organs has been cal role is suggested. shown to correspond with that of the III. Hormonal Control of Reproduction hermaphrodite gland (Runham and Laryea, 1968; Takeda et al,, 1986). In A. General Morpholegy of the Repro- some speeies, these hormones are steroid ductive System in nature. However, the ehemieal nature The reproductive system consists of has been still unknown in most other the hermaphrodite gland, the herma- species, Gottfried and Dorfman (1970a, phrodite duct, the albumen gland, the b) were the first to show that sex steroid common duct (plus free oviduct), the hormones such as 11-ketotestosterone and spermatheca and the penis together with dehydroepiandrosterone are synthesized vas deferens (Fig, 4). The rnorphology by the hermaphrodite gland in the and physielogy of the aceessory sex to slug, Ariotimax californicus. In Ezchadra organs differ somewhat from speeies peliomphala and Limax 7narginatus, speeies. In the infantile stage uf peripheral regions of the acini in the Euhadra pelio7npahala, the reproduetive hermaphrodite gland are shown to give a system consists of a simple elongated positive reaetion for 3B-hydroxy steroid tube, the proximal end of which is rudimentary herma- dehydrogenase. Furthermore, sex steroid eonneeted to the is hormones such as progesterone, testo- phrodite gland, and the distal part sterone and estradiol were detected in conneeted to the small penial sheath, organs are still the hermaphrodite gland (Takeda, 1980, The other accessory sex During 1983a, 1985a). The amount of sex steroid rudimentary or entirely lacking. shell, the reprodue- hormones in the hermaphrodite gland the growth of the small. With the of Euhadra peliomphala is sumrnerized tive system remains in Table 1. In the hemolymph of Li?nax development ef the reversed periosteme 1982b), the of the marg･inatzas, sex steroid hormones have (Takeda, growth NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda :Hormonal Control ofReproduction in Land Snails 107 A B C DA fN 1･1 rm 11LT 8 ' gjeAeqlt 213 "11l)sktw9 / 4 st' i Lglii/fgwo11 g? g12.e-12 ?5A -13 Fig. 4. Differentiation of the reproductive system in the snail, Euhadra Peliomphala. 1. penial sheath, 2. flagellum, 3. epiphallus, 4. vas deferens, 5. receptaculum seminis, 6,harmaphrodite duct, 7. hermaphrodite gland, 8. dart sac, 9, mucousgland, 10, vagina, 11, oviduct. 12, com- -t mon duct, 13, albumen gland. q K s･) d ?d ll ket 6 th reeefi'xopa･itomee. 1. wagng, 2. va)tkee, 3. kg, 4. thager, 5, ecreue, 6. detrrr, 7. pttrwa, 8, 9ege, 9, xavawa, 10. ve, 11. uatyNcr, 12, dittMut, 13. fieza. reproduetive system is rapidly com- (Takeda, 1982c). The gonacl consists of pleted. The abrupt growth and differen- a number of sac-like acini in which both tiation of the reproductive system is gametes mature. The maturation pro- related to the rnaturation of the gonad. cesses of the hermaphrodite gland in Limax morginatus are shown in Fig. 5. B. Maturation of the Reproductive Maturation of spermatocytes is fairly System rapid, so that although oocytes differen- a. Gametogenesis tiate first they reach maturity atter The hermaphrodite gland in snails spermatogenesis is largely complete. resembles bunches of grapes and in This type of maturation is termed pro- partially embedded in the hepate- tandrie hermaphroditism and is generally pancreas, this is different from that of found in slugs. The relative proportion slugs where a separate mass and arrangement of varied with it is garnetes / NII-Electronic Library Service The malaoologioalmalacological societysooiety of Japan 108 VENUS ： Vol ，48， No ．2 （1989） Fig．5． Stages in maturation of the hermaphrodite gland in the slug ， Limur marginatus ． − A ： undifferentiated stage ， B ： male phase ， C ： male female phase ， D ： female phase チ コ ウ ナ メ ク ジ の の A includingatrophy stage 、Scalet 50 μm ． ャ ラ 両 性 腺 成 熟段 階 ， ． 一 未 分化 期 ，B ．雄性 相 ， C ．雄 性 雌性 楓 D ，雌 性 相 （退縮 期 を 含 む ）． 一 NII-ElectronicN 工 工 Eleotronio Library Service The malacologicalsocietymalacological society of Japan Takeda: Hormonal Control of Reproduction in Land Snails 109 the progress of development, On the moval snails. On the other hand, ten- other hand, spermatogenesis and oogen- tacle removal snails into which extraet esis progress almost at the same rate in of the optie tentacle without the eptic snails. This type is called simultaneous gland was injected eontinued low game- herrnaphroditism. In both types, oocytes togenie aetivity. In eultivations ef the and spermatocytes have been known to hermaphrodite gland in a medium con- differentiate from the germinal epithe- taining optic glande extract, oogenesis lium under the influenee of hormones. did not proceed but spermatogenesis Experimental investigations of the regu- was promoted. In contrast, in the lation of gametogenesis have been medium containing extract of the optic undertaken with these speeies (Taka- tentacles, frem which the optie gland yanagi and Takeda, 1985>. In Euhadra had been removed, gametogenesis stop- peliomphala･, from which the optic ten- ped. Upon injeetion of brain extract taeles had been removed, both spermato- late ooeytes･ disappeared from the aeini genesis and oogenesis were inhibited. whereas early oecytes remained. Sper- This inhibition was reeovered by injeet- matogenesis was not influeneed. In the ing optic tentacle extract. Spermato- in vive system, brain extraet had a genesis in tentacle-removed snails whieh stimulatory effeet on oogenesis, but no received optic tentacle extraet was rnore effeet on spermatogenesis, After injec- promoted than that of the tentacle re- tion of testosterone, the number of male moval group witheut injections. In the germ cells inereased and this effeet was female part, further reductions in oogen- inhibited by anti-androgen. On the esis were noted when the optic gland contrary, after injection of estradiol extract was injeeted into tentaele re- the number of ooeytes inereased and ' Table 2. Effects of hormones on gametogenesls in vitro in the snail, Euhadra -?-t ' PetiomPhala. E JK ); ?-f oMma[]iJreanc ts m esi-e. rk ,) =e vozaasXec tset6njbl A: Spermatogenesis . ". . -.. .- a6'ptic"Eiana' b Germ Cells ontrol HormoneControlTestosteroneTes,+AA560 .C. . Spermatocyte 22.5 ± 1,8 19, 3 ± 1. 1 32. 7 ± 3. 5 20. 2 ± 6. 2 24. 0± 7. 1 Spermatid 20.0± 1.7 13. 3± 1, O 56. 3 ± 2. 9 52. 0± 6. 3 63. 0 ± 5, 5 Spermatozoon-i7. 2. 1 5± 67.4± 1,6** 11,O± 3.5 2Z 8± 4.9* 13.0± L5 B : ... Ppggpesis a b Germ Cells Control Brain Hormone Control Estradiol Est.+AE 10275 Early Oocyte 39,7 ± 1.7 28. 3± 3, 5 32. 7± 3. 5 2o. e± o. 6 45. 0± O. 6 Middle Oocyte 5LO ± L2 55. 4± 3, 1 58. 0± 2. 9 65,O± 1,7 46. 2± 2. 7 Late Oocyte 9.3 ± 1.5 16, 3 ± 2. 7** 9. 3 ± 3, 5 15, O ± 1, 2* 8.7 ± 2.3 Cultures were maintained at 23eC for 10 days. The total number of each type of germ cell per acinus is expressed as a percentage. Concentration of hormones: 1. 5 optic glands ,'ml medium, 1,5brains,/ml medium, steroid hormones 3ptg/ml medium, inhibitors 1 ptg/ 'p NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 110 VENUS: VoL 48,No, 2 (1989) their development progressed. When Lane (1961) have been applied by a pieces of the undifferentiated herma- number of investigators to various spe- phrodite gland were eultured in the eies of land slugs and snails, In Arioli- medium containing testosterone, ac- max cali.for7zicus, on the eontrary, optie celerated spermatogenesis was induced, tentacle removal in immature stages while the presence of estradiol pro- indueed preeoeious spermatogenesis and moted oogenesis. These effects were in- the injeetion of eptic tentacle homoge- hibited by anti-steroid substanees. These nate halted it (Gottfried and Dorfman, are summarized in Table 2 (Takeda, 1970a,b). Similar results were ebtained 1983a). by administering dehydroepiandrosterone The aetion of sex steroid hormones and 11-ketotestosterone to slugs with on gametogenesis has been examined by and without the optie tentacles, From gonadal regeneration in Mmax ?nargi- these results, it was suggested that na･tus. The epithelial cells of the cut spermatogenesis is eontrolled partially surface dedifferentiated, proliferated and by the eptic tentaeles and steroid hor- formed the regeneration bud. In Limax mones infiuenee the effect of the ten- margixatzas injeeted estradiol, ooeytes tacular regulator or modifier of gameto- were highly recognized in the regener- genesis, In Arion subfusct{s, a clear ated acinus and spermatoeytes were ob- feminization of the gonad was observed served with testosterone. Therefore, after removal of the optic tentacles at gametogenesis is shown to be controlled birth, while spermatogenpsis appeared by sex steroid hormones direetly (Ta- to be blaek in the operated slugs, This keda and Sugiyama, 1984). indieated that the optic tentaeles ex- Most of the researehes on the horrnonal erted a stimulating influence on sper- control of gametegenesis have been per- matogenesis and an inhibitory influence fromed with slugs by using classical on oogenesis experimental morphologieal techniques, These conclusions are in line with the Pelluet and Lane {1961) were the first original experiments of Pelluet and Lane to show the hormonal eontrol ef game- (1961), tegenesis in the slugs, Arion ater and In addition, the oeeyte growth has Arion sztbfuscus. They extirpated the been shown te be under the eontrol of optic tentacles and then injected homo- the dorsal bodies dispersed in the con- genates of the optic tentaeles and cere- nective tissue surrounding the cerebral bral ganglia, They eoneluded that the ganglia in Peroceras ret･ieulatz{m (Wij- optic tentaeles produee a hormone stimu- denes and Runham, 1977) and Arion lating spermatogenesis and suppressing subfitsczts eogenesis, and that the brain produce a hand, in the European snails, the effeets hormone stimulating oogenesis. The of tentaeular removal are net as dis- latter reaches its full potency at sexual tinct as in other speeies. In HeVix maturity. Therefere, gametogenesis ap- pomatia, removal of the optic tentaeles pears to be controlled by antagonistie redueed the gonial divisions and caused hormones eriginating from the optie oocytes to degenerate before maturity, tentacles and eerebral ganglia. The ex- but did not affeet oogenesis (Sanchez perimental proeedures of Pelluet and and Sablier, 1962; KUhlmann and Nolte, NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda: Hormonal Control of Reproduction in Land Snails 111 1967). When hermaphrodite gland ex- the souree of the masculinizing factor traet was injected into the tentaele re- is unknown. In Helix apersa, oogenesis moved snails, the number of oocytes and spermatogenesis still proeeed nor- increased (Bierbauer, 1978). In Helix mally even though the optic tentaeles aspersa, oocytes in the undifferentiated were removed for one year (Bride and rudiment ot the hermaphrodite gland Griffond, 1981). Thus, results obtained have been shown to undergo autodif- up to now are somewhat contradietory. ferentiation when the gland was cultured This may be the results of different ex- in the synthetic medium without hor- perimental eonditions or species spe- mones (Guyard, 1971; Gomot, 1973; eifieity. Gomot et al,, 1980), However, vitello- b. Aecessory sex organs genesis did not advance. In organ cul- Abeloos (1943) castrated various spe- tures, the cerebral ganglia stimulate the cies of slugs and observed a cessation growth of ooeytes. This growth is of the growth and a regression in size mainly due to vitellogenesis. In these of the reproductive tract. Laviolette snails, however, the dersal body hor- (1954) transplanted juvenile gonads mone has not been examined. Sper- and/er reproduetive traet into the hemo- matogenesis is dependent on the pre- coel of normal or of eastrated slugs, sence of a hormonal androgenic factor Maturation of the albumen gland and produeed by the eerebral ganglia. The the eomnion duet was elosely related to 'that tentaeles also stimulate mitotic activity of the hermaphrodite gland (Run- of the spermatogonial cells and inhibit ham and Laryea, 1968). In Ezthadra the differentiation of ooeytes. However, peliomphala, the development of the Fig. 6.Effects of castration on the development of the re- productive system in the snail. Euhadra PetiemPhala. Accessory sex organs undergo drastic regressive changes 30 days after castration. AG: albumen gland, CD: com- mon duct. :･ -9?l rti otsvaXirXo\iftr=ts lkea'-g-it bloblM(E : Xtre..Jts : ±es). AG: afewa,CD: madtthig. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 112 VENUS: Vol. 48, No. 2 (1989) aeeessory sex organs, sueh as the head- far more conspieuous than the other wart (Takeda, 1980), albumen gland, two. In Euhadra peliomphala, the epi- eommon duct and vagina were inhibited thelial cells of the head-wart underwent and atrophied after castration (Fig, 6), regressive changes following castration. However, after injection of hermaphro- However, with the injection of herma- dite gland homogenate or sex steroid phrodite gland homogenate, the head- hormones, these organs return to normal, wart recovered completely to appear Thus, the growth and differentiation of normal. The swelling and development the hermaphrodite duct and of the of the head-wart is shown to be related albumen gland are suggested to be cen- intimately to the function of the herma- trolled by sex stereid hormones or other phrodite gland. When testosterone was hormones produeed by the hermaphro- injeeted into the body cavity of castrated dite gland. snails, the head-wart developed gradu- 1. Head-warts ally and recovered from eastration. In 1935, Taki reported an irregular When the immature head-wart was polygonal tissue situated between the cultured in the medium containing two optic tentacles on the head of many hermaphrodite gland homogenate, it de- land snails (Fig, 7). He named it the veloped enormously. The epithelial cells "head-wart". There are three types of of the head-wart cultured with testo- the head-wart. The first type is a sterone developed sirriilarly. However, "head creseentic depression, called the no effect was found in the head-wart hollow"; it oecurs in the snails, Satsuma cultured with estradiol. The develop- and Chtorites. The seeond type, S-shaped ment of the head-wart was reflected not tubereles, appeared in the snails, Aegista only in the nuclear volume but also in and Trishoplita. The third type consists the elongation of the epithelial eells of a row or small mass of warty tuber- (Takeda, 1980). The development of the cles and oceurs in the snails, Euhadra, head-wart is shown to be eontrolled by Bradybaena, Dolicheuota, and Heli- testosterone secreted from the herma- costyla. Among theTn third type was phTodite gland of snails. As described Fig.7. Swollen head-wart (arrew) in the snail, -vl Euhadra PeliomPhala. i - l "d op twltLkutbe (eE:a]), NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda; Hormonal Contrel of Reproduction in Land Snails I13 later, structures related to the head- (van Minnen and Sokolove, 1984). A wart have been repoTted in some speeies similar molecular weight range has been (Binder, 1965, 1977; Chase and Michele, found in Helix pomatia (Goudsmit and 1981). The develepment of these struc- Ram, 1982). On the other hand, in the tures is also suggested to be under the slug, Astliol･iwzax colu7nbianus (Meenak- hormonal eontrol of the gonad. shi and Scheer, 1969), the incorperation 2. Albumen glands of glueose Ci` into galaetogenin of the The albumen gland is yellow or trans- albumen gland is greater following optie parent and is situated against the an- tentacular remeval than in controls and teriDr coneave surfaee of the digestive this effeet is reversed by the injection gland, It is a compound tubular strua- of optie tentaeular extract. In Achatina ture eonsisting of seeretory cells inter- fulica, the dorsal body is developed spersed with ciliated cells and it seeretes enormously. When pieees of the un- the granules of galaetogen produced by differentiated albumen gland were eul- the Go!gi appaTatus smit, 1969), Galactogen surrounds eaeh body homogenate, the gland became fertilized egg and is catabolized by the mature. However, it did not differentiate developing ernbryo. The albumen gland in the medium eontaining brain homo- fluetuates in size and galactogen eon- genate without the dorsal body (Takeda tent during the reproductive eycle: it is and Takeda, 1985), In Helix aspersa, largest and the galactogen content high- injeeted optie tentaele homogenates have est during the breeding season and is been shown to eause a deerease in the minute and the content lowest during growth rate and the albumen gland the period following egg laying. Matu- development (Bride et al., 1986). ration of the albumen gland has been In Limax marginatus, the presenee of shown to be under the endoerine eontrol, a gonad is required for the growth and The brain of Helix pomabia･ produces a differentiation of the aceessory sex neurohormone called galactogenin, which organs including the albumen gland stimulates galactogen synthesis in the (Takeda et al., 1986). In Euhadra albumen gland trary, the major souree of the galactogen has been shown to be eontrolled by a sex synthesis-stimulating faetor has been steroid hormone, estradiol, seereted by reported to be the dorsal bedies (van the hermaphrodite gland (Takeda, Minnen and Sokolove, 1984). This was 1983a>. Thus, the hormonal control of also shown by removal ef the dorsal maturation of the albumen gland is bodies in DeroceTas reticulatum (Wij- highly speeies specifie. denes and Runham, 1976), The aetive 3. Reprod-u.etive tracts princip]e in Limax maxi7n2ts does not The lurnen of the common duct is stimulate galaetogen synthesis in Helix partially subdivided by lateral folds into pomatia and vice versQ (yan Minnen the male and female ducts. The gland et al., 1983), Fraetionatien of whole opening into the male duet eenstitutes brains in bima･x maximus yielded a the prostate gland and those opening molecular weight range of 4000-7000 into the female duet the oviducal gland, NII-Electronic Library Service The malacologicalsocletyofmalacological society of Japan 114 VENUS: VoL 48, No. 2 (1989) The prostate gland of most stylommato- the transition period, both the male and phoran species consists of thin, long female parts of the implants developed. tubular glands conneeted to the sperm Therefore, two hormones are involved groove along much of the length of the in the growth and differentiation of the spermoviduct. Most prostate tubles have common duct, eontrolling the male and two types of cells, gland eells and female parts of the tract, respectively. ciliated non-secretory cells. The female Bailey (1973) has shown that the gonad part of the spermoviduct is extremely fails to stimulate the development of glandulaT. The duct wall eonsists of an the rudiment of the reproductive tract inner eiliated epithelium, the middle of Agriolimax reticztlat?ts in vitro. How- glandular layer and the outer connective ever, the rudiment starts to develop when tissue. Runham et al, (1973) have ex- the gonad, the juvenile tract and a brain tended Laviolette's results on Deroceras are present in one culture medium. How- retietttatum and showed that implanta- ever, it is not reasonab]e to eonsider that tion of undifferentiated tracts into slugs the gonad stimulates the brain to pro- in the early protandric phase resulted duee a male and female faetor. As dorsal in the growth and differentiation of the body removal induces the atrophy of the prostate glands, whereas the fernale ovidueal gland, development of this parts of the implants remained undif- organ is suggested to be under the hor- ferentiated and small. Transplantation monal control of the dorsal bodies (Wij- of the reproduetive tracts of juvenile denes and Runham, 1976). slugs into the hemoeoel of egg-laying In Limax ?itaTginatus, eastration also slugs was followed by the exclusive de- led to the atrophy of the accessory sex velopment of the oviducal gland. During organs. When the castrated snails were =-･ {{g '" ' 'lil'1 .", ,I7,,- ,I.,"t t//{tt'liii'm,,:'..f･l 1'i,l. , ,illil ,ms.:. '-'' fi .wt ltt- ='f,uil/ tw.I' '.tt ., '/vt/ ttt,II' . . ltl . , , . ' ' i" /t G i'' , 1 i ' ' 1://ttttt ,r . - 'L'''ma"-diti'nve E/ ,'f'. t, ,//, . f/ ,l . #,, .,, , A B.' . ,li･ie'. :,llll,iil'K- ,um.･ Fig. 8. Effects of sex steroid hormones on differentiation of the common duct in the slug, Limax marginatecs. A: control, B: te$testerone treated, C: estradiol treated. Pieces of undifferentiated common duct were cultured in a rnedium containing steroid hormones for 10 days at 23eC. OG: oviducal gland, PG; prostate gland, O: oviduct, DV: vas deferens. Scale 30 ptm. f -v = V it] fi yamb{thwheroG>TL IC ts lk V:'t-7" I F' rk ,VtvObl ee . OG: ravaua, PG: ditri sZwa, O: thgNff, VD; wtreee. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda: Hormenal Control of Reproduction in Land Snails 115 injeeted with hermaphrodite gland homo- castrated snails into normal adults in- genate, the common duet including the itiates seeretory aetivity. These experi- prostate and oviducal glands enlarged. ments show that the presenee of the 'testo- Jn vivo experiments showed that gonad is essential for the multifid gland 'the sterone was shown to affect develop- to develop seeretory activity and that ment of the prostate glands and estradiol the influence is mediated by an endocrine of the ovidueal gland (Takeda et al., factor, In organ ¢ ultures, the inactive 1986) (Fig. 8). glands are aetivated rapidly when a In Euhadra petiomphala, effects of pieee of the gonad is added. This activa- eastration were morphelogically unelear. tion is chaTacterized by the increase in However, by the injeetion of the herma- synthetic aetivity of the cell organelles, phrodite gland extraet or estradiol, the The stimulus ef the nervous systern re- vaginal epithelial cells of juvenile snails sults in a release of secretory substanees were shown te inerease their length. into the ducts by the gland cells. The 'to The penis is a highly muscular tissue. multified gland in Helix eoTresponds In Deroceras reticu,latu7n, the degree of the mueous gland in Euhadra. As shown the growth and differentiation of the in Fig. 6, no morphologieal changes immature penial eomplex implanted into were observed following castration. the hemocoel of host slugs depended en The Telationship between the brain the initial stage of host and implant hormone, the gonadal hormones inelud- (Wijdenes, 1981), Implants never de- ing non-stereidal materials and the veloped in castrated slugs and was net dorsal bedy hormone on maturation ef under the eontrol of the grewth hormone the reproduetive system require more from the brain (Wijdenes and Runham, detailed studies. 1977). In this speeies, it is suggested C. Mating Behayior that･ the gonad produees a rnale hor- mone. Furthermore, as the growth of Although snails are typieally herma- the penis was shown to oecur in im- phrodite, some of whieh can propagate mature slugs implanted with the mature by self-fertilization, they still seek other gonad in Liwtax maximtts, the existence snails fer eopulation and hence exehange of gonadal hormones is presumed and of sperni (Takeda, 1983b, 1985b). In neurohormones seem to act indirectly the adult sexually mature snail, Et{･hadra via the gonad to promot the developrnent peliomphala, the head-wart attains its of the reproduetive traet (MaCrone and full size and t.he penial mass is distinct Sokolove, 1979, 1986; Sokolove et al., and sometimes slightly protruded. When 1984, 1985). the snails meet in the breeding season, Gomot et al. (1980) and Gomot and they begin to touch the partner's head- Colard (1985) have shown that the wart whieh then swells gradually and gonad and the eentral nervous system forms a conspieuous hemispherieal pro- haye a stimulating influence on the trusion (Fig. 7). The swelling and con- multificl gland. In Hel'ix aspersa, after traction of the head-wart is repeated eastration the glands deerease in ac- several times. Two snails may remain tivity. Transplantation of non-differen- head to head for a long time, Then the tiated eells from the multifid gland of head-wart gradually eentracts as the NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 116 VENUS: VoL 48,No. 2 (1989) penial mass begins to swell and the the head-wart cells may be modified frorn snails begin to copulate. Similar sexual ordinary epithelial eells. behavior ean be experimentally indueed It is well known that the inseet sex in sexually mature snails by the use of pheremone shews a strong attractant ef- head-wart extraet. In juvenile snails feet. On the other hand, the snail sex outside the breeding season, prominent pheromone is more aphrodisiac than activity did not appear. In castrated attraetant. These differences are also snails, no sexual excitement is observed, seen in the fine strueture of the gland The aetive principle released from the ee]ls (Takeda, 1983d>. DiffeTent from head-wart is a kind ef the sex phero- inseets, the sex pheromone in snails is mones which induces sexual excitement mainly synthesized by the Golgi ap- (Takeda and Tsuruoka, 1979>. The sex paratus and may be proteinaceous in pheromone is released to the sexual nature. partner frorn the microvilli by protrud- The most eomplete study of reproduc- ing their head-wart (Takeda, 1983d). tive behavior has been made in Hetix This pherorr}one does not appear to serve pomatia (Jeppesen, 1976, Lind, 1976) as a sexual attTactant, but rather to and Helix aspersa (Meisenheimer, 1912; sexually excite another snail, thus evok- Herzberg and Herzberg, 1962), The ing a eourtship response following their four stages in matang behavior were initial eneounteT. Similar speeialized deseribed. These are the spiral upright dermal struetures are present in the Te- phase (eopulation oeeurs during this gion between the optic tentacles in some stage), the upright phase (sexually re- other land pulmonates, These structures sponsive snails meet head-on and re- have also found to have a funetion in peatedly touch the tentacles), the down courtship behavior. These are the turning phase (the love dart is foreed "Cephalic dimple" of Achatina aehatina･ out into the partner) and the imrnobility "Frontal (Chase and Michele, 1981), the phase (after copulation, both partners organ" of the snail, Gymnario7i (Binder, remain immobile), In about 10 families 1965, 1977). The frontal organ is situ- of land snails, the existence of daTts has ated above the snail's mouth, between been known and the major funetion is the four tentaeles and usually rather possibly associated with reproduetion, nearer the dorsal pair. Furthermore, The dart is extruded through the genital "Fossette the triangulaire causale" pore and pushed into the mating part- "Caudal in the dersal rnid-line and has some that the dart glands show activity not similarity to the cephalie dimple. These only during eopulation but also during dermal structures are not pathological egg-laying. The development of the dart but normal and represent some specific gland is suggested to be controlled by funetion in eourteship behavior, except the gonad the head-wart of the snail, R?t?nina de- snails are not only able to eopulate eollata (Miles, 1961). In pulmonates, sueeessfully but also actually have a all the epidermal cells haye a microvillar slightly higher rate of successful copula- structure (Wondrak, 1967). Therefore, tion than do eontrols. Many elaborated NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda: Hormonal Control of Reproduction in Land Snails 117 courtship sequences have been deseribed tend to increase their behavioral aetivity in s]ugs. In Li7nax maximus, the spee- in rainy or moist conditions, Environ- taeular aerial mating, hanging in pairs mental water seems to play a dorninant by thiek mucus repes suspended from role in aetivity of terrestrial pulmonates. trees has been reported (e.g. Fischer, Bahavior of terrestrial pulmonates is 1917). In Arionid slugs, the caudal largely divided into two types; one is gland only reaches the maximal develop- eircadian loeomotor behavior and the ment with sexual maturity It is an indicator of the degree of sexual behavior maturity, suggesting the control by Takeda, 1987b). In both cases, feeding gonadal hormones. Courtship behavior behavior and reproductive behavior are also seems to be under the control of usually observed. Organs of land pul- gonadal hermones, because it is in- monates also increase in water content hibited by eastration. with the progress of hydration, and viee Although the eourtship process is in- versa･. Induction of loeomotor bahavior dueed by the sex pheromone released is considered to be rnediated by the from the head-wart, activity of snails nervous system. The nervous system in the breeding season is highly ac- bathed in the hemolyrnph is also in- eelerated by rain. Many observers have fluenced by hemolymph osmolarity. In noted that rain brings about inereased Achatina .fulica, effeets of osmotic stress mating. In general, land pulmonates on electrieal aetivity have been examined WATERENVIRONMENT 'i iumidit.y.. E.y'dr'Eti.otn.Ri?ls/c.al`p.n]': HEMOLYMPH eSMOLALITY - Hypertonic Hypotonic g CENTRAL NERVOUS SYSTEMT -# Active lnactLve BEHAVIOR Active - tnactive e m)t--ooreso-><=wom!-o"[t-- 6 o circadian Rhythm l ] Active a' en - o.- t-t $ rnactive W L 130 230250 300 X Kg H2o } Hemolymph osmglZ]Ritsym Fig. 9. Schematic drawing of the induction of locomotor behavior in land pulmonates. wntiIRifVLtsO-6fiuteeitCDeeifblhkS-ma rtpa. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 118 VENUS: VoL 48,Ne. 2 (1989) in vitro by changing the bathing rne- D. Oviposition dium. In the nerve eells the parietal In slugs, ovipositien has been known ganglion, electrical aetivity was indueed to be under the hormonal control (Ta- by hypotonic saline and suppressed by lceda, 1977, 1979). In Limax flavus and hypertonic saline (Takeda and Hase- Deroee'ras reticulatum, brain extract gawa, 1984; Hasegawa and Takeda, during the breeding season stimulates 1986), In Deroeera.s reticttla･tu7n･, rates egg-laying and optie tentacular extract of eleetrical diseharges in the pedal inhibits it. The rate of fertilization in ganglion increased as the bathing fiuid slugs injected tentaeular homogenate beeame more dilute (Hughes and Kerkut, was lower than that in those injected 1956; Kerkut and Taylor, 1956). brain homogenate. Removal of the optic Furthermore, changes in the osmotie tentacles induced a large inerease in the pressure of salines ean cause rapid numbeT of eggs laid and an increase in changes in specific nerve eells in the the rate of fertilizatiDn in both species. isolated brain of the slugs, Limam pse7Ldo- Aceelerated oviposition and a high rate flavzts and Li?nax maxi7nus (Prior, 1981, of fertilization indueed by removal of 1985). the optic tentaeles were inhibited by the Therefore, it is reasonable to eonsider injeetion of tentaeular homogenate, t･hat some nerve cells are involved with Furthermore, egg-laying was shewn to the induetion of loeomotor behavior be stimulated by estrogens such as when hemolymph osmolality was redueed. estradiol and estrone. On the other hand, The meehanisms of the induction of androgen treatments with testosterone loeomotor behavier in terrestTial pul- and dehydroepiandrosterone did not so monates are proposed sehematieally affeet the number of eggs oviposited, (Fig. 9>, Circadian behavior is induced However, the rate of development in by conditions of relative humidity of this group was comparatively higher over 5092 and of hemolymph osmolality than that in estrogen groups. Estrogen from about 120 to 230 mOsm. Endo- treatments increase the number of eggs geneous noeturnal rhythmicity is under oviposited and androgen treatment en- the control of the eentral nervous system. hances the rate of development speeifi- As to hydration-related behavior, at first eally. In a joint injection with de- water penetrates the body cavity through hydroepiandrosterone and estradiol, the integument and reduced hernolymph larger number of eggs were oviposited osmolality. Then, some nerve eells in and the rate of development also in- the centra] nervous system are activated creased, Sinee the gonadotrophie hor- by the osmotic stress and induce loeo- mone inereased egg-laying', and inhibi- motor behavior. In contrast, in dry con- tors redueed both the rate of egg-laying ditions, evaporation of water through and develepment, regulation by steroid the integument leads to an inerease of hormones of oviposition is suggested. hemolymph osmelality, The central It may be possible that the faetor nervous system is inaetivated by the directly coneerned with egg-Iaying is osmotic stress and Ioeomotor behavior not the brain hormone but steroid hor- stops, mones in these speeies. A brain hor- mone, therefore, seems to act as a NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda: Hormonal Control of Reproduction in Land Snails 119 gonadotrophie hormone, stimulating the gametogenesis. If food is abundant, gonad to secrete estrogens. As described individuals attain reproductive size later, neurohormones have been shown more rapidly and produce mere gametes, to induee oviposition in Aplysia and In invertebrates, sex steroid hor- Ly'm?taea. mones have been detected and t･heir physiological roles have been examined IV. Dynamics of Hormonal Control in (Sander and Medhi, 1979; Sandor, 1980). In the present speeies, sex Reproduetioll steroid hormones are shown to be in- To propagate the speeies in a severe velved in the control of reproductive land habitat, hermaphroditism is a very aetivity in a way surprisingly similar useful method for slowly moving snails to that in vertebrates. In other speeies, with low density populations. Herma- such as Helix potozatia, histologieal phroditism has generally been thought ehanges in the reproduetive system to represent a primitive pattern of have been examined by treating with sexuality in molluscs (Ghieselin, 1969). steroid hormones: testosterone stimulates Environmental faetors whieh affect the the male phase and inhibits the female development and maturity of the repro- phase, estradiol has the reverse effeet, ductive system are temperature, rnois- while progesterone stimulates both ture and photoperiod (Bouillon, 1956; phases (Aubry, 1961). Wolda, 1967; MeCrone et aL, 1981). On the basis of the results mentioned These faetors finally affeet the endo- above, a tentative hypothesis is sug- erine system and induee the secretion of gested for the endoerine control of hormones (Geraerts, 1986; Gomot, hermaphroditism in the present land 1986). In maturation of the reproduc- pulrnonates (Fig. 10). Male and ferriale tive system of slugs, Deroceras reti- aetivities are controlled by different hor- culatum and Arion a･te7', humidity and mones. An androgenic neurohormone is light exert a greateT effect en sperma- produced firstly by the brain. This fae- togenesis than on oogenesis (Lusis, tor stimulates the optie gland to seerete 1966; Smith, 1966; Runham and Layea, a hormone, On the centrary, female ae- 1968). In Limax maximus, photopeTiod tivity is controlled by a neurohormone is critical in male phase maturation but produeed by the brain. When the her- has Iittle effeet on female phase matura- maphrodite glands were eultured with tion (Sokolove and McCrone, 1978). the brain and optie gland, seeretions d'f Sexual maturation can be postponed by sex steroid hormones were facilitated. keeping the juvenile slugs at short day Testosterone secretion was stimulated photoperied, and exposure of the im- by the optic galnd and estradiol secretion mature slug's to a long day photoperiod by the brain. Therefore, both the brain induees sexual maturation and MeCrone, 1978; McCrone and gested to aet on the hermaphredite gland Sokolove, 1979). In addition tQ these as a gonadotrophie hormone. In Dero- factors, food may effect the timing of ee･ras ret･ict{latu7n, the injection of pitui- reproductive onset in individuals via tary hormones acce]erates the growth its influenee on the growth and rate of and maturation of the gonad (BTidgeford NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 120 VENUS: Vol. 48, No. 2 (1989) LIGHT FEMALE PART -er MALE PART T t t opTtCG+/Ltt>AND e + HERMAPHFrc}DITE GLAND Fig. 10. Schematic drawing of the hormonal control of reproduction in land pulmo- nates,ma{liHmalttsV'r6LkfuX'k'XOD' ft ･ rNnto rk ?v - v fi ee Ef fifttgejS mu. and Pelluet, 1952). Among sex steroid as the third order have only been demon- hormones, testosterone stimulates not strated in vertebrates, In vertebrates, only the differentiation of the rnale cells the eontrol of repraduetion is achieved but also the growth and differentiation by the hypophysis and the gonad inter- of the male part of the reproductive ealates between the hypothalamic neuro- tract. Estradiol eontrels the growth and hormones and the aecessory sex organs, differentiation of the female part of the It is interesting that the dorsal body of reproductive tract. In some species Helix pomatia has a capacity to syn- whieh have developed dorsal bodies, the thesize steroid (Krush et al., 1979). The dorsal body hormone stimulates the de- relationship between the gonad, the brain velopment of the female aceessory sex and the optie gland was examined in organs and maturation of oocytes. The E?thadra peliomphala (Takayanagi and relationship between the dorsal body hor- Takeda, 1985). The nuclear volume of mone and sex steroid hormones in repro- the optic gland eells cultured in the duction are the problems to be resolved, medium containing hermaphrodite gland Thus, male and female activities are extraet decreased eompared to controls. controlled via first. second and third However, the volume did not ehanged order neuroendocrine mechanisms, as signifieantly in the medium containing described later. Such higher mechanisms brain extract. On the other hand, the NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda: Hermonal Control of Reproduction in Land Snails 121 nuclear volume of the optie gland cells into seven elasses: Polyplaeophora, also decreased in the medium with testo- Aplaeophora, Monoplacophora, Scapho- sterone. However, the volume did not poda, Bivalvia, Gastropoda and Cepha- ehange in the medium with estradiol, lopoda. Among these, the endocrine een- This suggests the presence of a feed- trol of reproduetion has mainly been baek system between the gonad-inhibitory studied in gastropods and cephalepods. hormone in the optie tentaeles and In other classes, researehes on reproduc- gonadal sex steroid hormones is also tien have simply been performed in the suggested in slugs (Gottfried and Dorf- level of environmental factors (Giese man, 1970a). Sueh feedbaek eontrols and Pearse, 1979). are well doeumented in the vertebrate Although rnueh has been written on endocrine system, An inereasing num- reproduetion of bivalves ber ef studies have dealt with the oceur- 1984; Lubet, 1986), researches on the rence and metabolism of steroids in hormone eontrorling sexual differentia- all land pulmonates. tion and gonadal activity are poor (Herlin-Houtteville and Lubet. 1975). V. Comparative Endocrimology of Neuroseeretory eells have been commonly Reproductian in Molluscs observed in the cerebropleural, viscera] In the phylogenetically lower species, and pedal ganglia. During the p.eriod as deseribed before, endoerine organs are of gamete maturation of Mytilus, the absent. In such species, neuroseeretory neurohormones aecumulate in neuroseere- cells of the central nervous system exert toTy cells of the cerebropleural and the direct eentrol over the target organs visceral ganglia (Lubet, 1955, 1956>. In and proeeed reproduction. This type is Dreisse"ta･ "zaxim?ts, diseharge ef neuro- ealled a first order endoerine meehanism, This is the most the intensive oocyte growth and spawn- fundainental mechanism and still pre- ing (Antheunisse, 1963), Thus, the sents in the other meehanisms, Then, a gametogenic cycle seems to be linked to second order (two step) neuroendoerine a neurosecretory eycle. However, gan- mechanism appears in the rnore highly glion removal experirnents eontradiet oTganized species where neureseeretory the relationship between neuroseeretion eells exert trophic effect on newly de- and the oocyte development and also veloped epithelial endocrine glands, hor- spawning. In the seallop. Patinopectetz mones from whieh subsequently affect yessoen,$is, spawning has been shown reproduetion. A third order (three te be induced by serotonin and be regu- step) neureendoerine meehanism is that lated by prostagrandin neurosecretory cells exert trophic effeets Nomura, 1987). Although the actions of on epithelial endocrine glands, hormones neurohormones are different from spe- from whieh are trophic to control ac- cies to speeies, regulation of reproduc- tivity of the next epithelial endoerine tion in bivalves is considered to be a glands on repToduction. Reproduetion fir$t order neuroendoerine mechanism. proeeeds more eMciency by the introdue- The class Gastropoda consists of three tion of feedback mechanisms. subelasses: Prosobranchia, Opistho- The Mollusea phylum is subdivided branehia and Pulmonata, In proso- NII-Electronic Library Service The malacologicalsocietymalacological society ofJapanof Japan 122 VENUS: Vol. 48, No. 2 (1989) branehs, reproduction has been examined Patella vulgata･ (Choguet, 1971), when in some species (Fretter, 1984), In the gonads are eultured with the cere- CTepidula fo7hnicata, which show sex bral ganglia or hemolymph of male reversal. the nervous system secretes a phase animals spermatogenesis ensues male factor for the male differentiation whereas, if the gonads are eultured during the male phase and secretes a with cerebral ganglia in the female female faetor during. sex inversion and phase they show aecelerated oogenesis. the female phase whieh initiates the In this c]ass, therefore, the control of female differentiation in the male phase reproduction is simply performed by a gonad searehes have been made on the hor- However, in some speeies of proso- monal centrel of penis differentiation branchs sueh as Gibbuta umbilicalis, and dedifferentiation (le Gall and Monodonta lineata (Clare, 1987; Herbert, Streiff, 1975; le Gall, 1981; le Gall et 1982) and Haliotis "efescens neuroseeretory eells of the right pedal ganglionar organ has been reported as ganglion aeeumulate in portal vessels of a putative endocrine gland. In Haliotis the right tentaele. This hormone stimu- rufescens, activities are shown to in- lates the differentiation of the penis in crease at the maturation period and to young males and disappears progres- deerease after spawning (Miller et al., sively in the female stage. The other 1973). However, details are unknown. neurohormone produced by neuroseere- In opisthebranchs, sueh as Aplysia, tory cells in the pleural ganglia aets on reproduetion has been examined (e.g, t･he penis and induees the reduetion of Hadfield and Switzer-Dunlop, 1984) and the penis. In Crepidula･ fornicata, the juxtaganglionar organ has also been Calupteo"a sinensis (Streiff, 1967) and found on the dorsal or ventral surface s lo lt. 20 ls 11 ls "cidtc Puptide/ S,e.lilklS'/:Iil'/1:II::i:'i!l'' aH/ ,'[il, .c.im±ts A"111t ::Ii !':1/ri,l'..ts.:i2':tth.-:i:-'.Si:I:::Ili,gl:..l-l'L;IT:e"-r･ysbi-iie-spr-tsn-Leupmm Lirge [ fi ;/ll#E･ige.g.lllii,iSIIi.i,,･iglJ-:-',,I:･i,,l.#lgll･li,',i:I,-.[,.He,ii･IIexIIEI･1ii,lfii;".=T,ii:fi･:ittg･1ac-#kagi-Bg,'L'{2gl,kEth･,X･l[-f["G,1,i,:I,A:r:Il･E:1,iEi,I:･IIImi･ss w{eA:}/ EUICAP1t lt:Il';il:I:PI:l:I::[:2;JLI[;[alnta"It/:!i't[hrk:IX[:.fijTtr:l:l4"l:iilg5}/:CEilir+'iulft:wt,u:lililil.l'.::!YatL/'#:I:::L']iting,:LLe"t:::tW"LYS't'i.ii / i " h W- : t'igE t,,.lei't.":lrii'ililli,iliiil,!fl,i,r'll'l "iE' Le.l,illii'ii" r "i"F'"'i /･IH : FepLiden: .S'e".. T'eptide B : ,tt,, t"i,il,ll'Eilllillll"=:t,,g/:hl,lll,ll,'ii.iiiZ#,ili'ii\,,i"..ll-ll,;g4ili,l,tL"l'iii/ilil,,-i-i:.gll,l$l,-s,tl Mi "i"e' X Rt-., / pgGLi{ajE}･-''･"M[fi] £mo,,.stE.'t,if7]] iile.ii;Gli.il"if "i'EFG"i'I.ii.Usii,iE,il':.;ri/l.I/Lllltr'5/1/bS"'Veiiiii,i/Iil'i'fL' ll?ttt,ll'ill' Fig. 11. Amino acid sequence homolegies among egg-laying related peptides in APlysia (AC: APlysia calijbrnica, AP: APlysiaParvula) and L,1,mnaea stag"alis, ELH: egg- laying hormone, ERH: egg-releasing hormone, CDCH: caudodorsal cell hormone (ovulation hormone), BCP: bag cell peptide, CDCP: caudodorsal cell peptide. Homologous regiens are enclosed wi'thin boxes. 7 1 7 i Yt -i 7 7 7 if'I OeegN caifK7'iF FuecD7 E 1meGdl)JEea)Nfii]tCi, ELH: NijNrii,v-r7, ERH: tyNtuffl;s 'f), ,v:e v, CDCH: KNrate;ti7v== 7 (VFij[Ufi:,v-e BCP: fiande-<7'ff- 1･', CDCP: -(i -(Ti5 eekb*Mve-< V' ,f N, NNMSG)}t 7 fi wn ,L 6. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Takeda: Hormonal Control of Reproduction in Land Snails 123 of the eerebral ganglia 1987). In other class polyplacophoran active peptides A and B with 34 amino amphineurans, similar organ, called the aeid sequence in eaeh. Peptide A and B ``Organ Juxtacommissural" have also eause egg-laying by activating the bag been present between the intereerebral cells to release the egg-laying hormone commissures and an anterior blood space (Heller et al., 1980). The egg-releasin.v. (Martoja, 1965; Vincent, 1970), Not hormone ean ulso act in this manner, only the juxtganglionar organ but also although it shares with the egg-laying the juxtacommissural organ has also hormone the capacity to stimulate egg- been thought to be a homologous organs laying in animals from which the bag te the dorsal bodies. Although some eells have been rernoved (Sehlesinger et studies have been perfoTmed on the al,, 1981), From the atrial gland, an- endocrine function ef the rhinophore in other peptides called ealifins whieh eon- AI)lysia (Vincent, 1969), there is no sists large three (A, B, C) subunits and information on the control of gameto- small subunits have been isolated and genesis and sexual maturation. In sequenced (Rothman et al., 1986), These Aplysi.a, researehes on the hormonal peptides show strong homology to the control of egg-laying have developed several bag cell peptides. These are surn- enormously (e.g. Geraerts et al,, 1988). marized in Fig. 11. At the present time, Egg-laying in Aplysin is regulated by researehes are developing to the gene the neurosecretory bag cells, loeated at level (e.g. Scheller et al.. 1984; Mahon the junetion of the parieto-visceral gan- et al., 1985; Nambu et al. 1986; Joosse, glion and the pleurovisceral connectives 1987; Geraerts et al,, 1988; Takeda, 1981). The amino acid sequenee of the mate homology of amino acid sequence bag cel] peptides, ealled the egg-laying between the egg-laying hormones and the hormones, was determined (Chiu et al., egg=releasing hormones. It is still un- 1979). It eonsists of 36 amino aeid elear, however, in what way the atrial residues. In addition, a mueh smaller gland peptides are involved in egg-laying. peptide, a-bag cell peptide (BCP) ean In this elass, the eontrol of reproduction also indueed the bag cell discharge is still perforrned basically by a first (Rothman et al., 1983>. Another pep- order neuroendoerine mechanism. tides have also been isolated from the The subclass Pulmonata is subdivided bag eells: these are acidic peptide, B, r into two orders. Basommatophera, and 6 bag eell peptides (Scheiler et al., largely Iiving in freshwater, and Stylom- 1983a,b; Rothman et al,, 1983, 1985; matophora mainly deseribed in this Mayeri and Rothman, 1985; Mayeri et review. Research on repreduction is al., 1985). Furthermore, the atrial gland highly advanced in Lynznaea stagnalis (Beard et al., 1982) in the reproduc- (Jeosse, 1975, 1979; GeTaerts and Joosse, tive system has also been shown to eon- 1984; Joosse, 1984, 1988), Female repro- tain three peptides that can induce egg- duetive activity is controlled by a neuro- laying when injeeted into mature ani- hormone and the dorsal body hormone. mals (Arch et al., 1978, 1980; Negel et The target for the dorsal body hormone al., 1985, 1986), These are ealled as the is the folliele eells and the hormone may NII-Electronic Library Service The malacological society of Japan 124 VENUS: VoL 48,No. 2 (1989) aet by activating the adenylate cyclic effect in maturing young snails (Ge- AMP system (de Jong-Brink et al., raerts, 1976). This is an example of a 1986). As a result, the dorsal body first order neuroendocrine mechanism. hormone stirnulates vitellogenesis in the In cephalopods, reproductive phene- eoeyte as well as growth and secretory mena have been reviewed aetivities of the female accessory sex 1976: Arnolds, 1984; Takeda, 1986>. In organs. It is the main female gonado- Octopus or Sepi,a, the central nervous trophic hormone. Maturation and ae- system has evolved to a complexity re- tivity of the dorsal bodies are contrelled miniscent of the vertebrate brain. Exten- by neurosecretory cells located in the sive fusion of the ganglia have altered lateral lobes (Geraerts, 1976), There- the situation ef the generalized nervous fore, this is a second order neuroendo- system ef molluscs, The Cephalopoda is crine meehanism, the first to be demon- independent of a neurosecretory common strated in this order. Although Sertoli pathway and utilizes the central nervous eells have been suggested to produee sex system to eontrel the production of the steroid hermones, steroids synthesized gonadotrophic hormone. This is per- in the gonad have simply a local hor- phaps due to the fact that the relation- monal effect (de John-Brink et al., 1981). ship between visual stimuli and optic In basommatophoran species, a feedback gland activky has beeome so elose. The effect from the gonad to the dorsal eontrol of reproduetion is comparatively bodies has not been dernonstrated (Boer simple, This may be due to the fact that et al., 1976). Egg-laying and egg-laying these species die soon after reproduc- behavior in Lymnaea stagnalis is in- tion. The only endocrine gland is the itiated and eoordinated by the ovulation optic gland (Wells, 1978>. The optic hormone seereted from the caudodorsal glands are small round bodies found on cells (CDC) Iocated adjaeent to the the upper posterior edge of the optic intercerebral commissure in the cerebral tract and are known to be of nervous ganglion (Gearerts and Bohken, 1976; origin et al,, 1985). Surprinsingly, the ovula- oka et al., 1970). The optic glands have tion hormone of Lymnaea is similar in an inhibitory nerve supply from the sub- amino acid sequence to the egg-Iaying pedunculate lobe of the brain. Therefore, hormone of Aplysia (Chiu et al., 1979). cutting its nerve conneetion results in A few specialized CDCs release a small an activation of the optic gland, fol- peptide whieh excites the other eells of lowed by a start of reproductive activity their network. a-eaudodorsal cell peptide (Wells and Wells, 1959). Sexual ma- (CDCP) has a positive feedback on CDC turity is regulated by seeretions of the firing activity whereas the B-CDCP also optic gland which are composed of the have a transmitter role, As to sper- stellate glandular ce]ls. The optic gland matogenesis, the small lateral lobes of hormone is not sex specific and it stimu- the eerebral ganglia have a stimulating lates the multiplication of the follicle NII-Electronic Library Service The malaoologioalmalacological societysooiety of Japan Takeda ： Hormonal Control of Reproduction in Land Snails 125 cell development the athird order neuroendocrine mechanism ． ， proteinaceous yolk − Like vertebrates the function of production and the growth of the ac ， gonadal − cessory sex organs in females ， and the homrones is to control gamete produc multiplication of spermatogonia the tion and accessory sex gland activities ． ， ， − production of spermatophores and the Fur 七hermore ， the control of the gonado growth Df the male ducts in males trophic centers is achieved by feedback − Wells and Wells 1975 Well 1976 mechanisms ． In these species ， the neuro （ ， ； ， ， 1978）．The chemical nature of the optic secretory and gametogenic cycles are gland hormone is sti11 unknown ． There dynamically linked through a series of ． have been no indications on the endo − feedback controls ． It is interesting to crine functien of the gonad ． However ， note that a third order neuroendocrine mechanism considered to the gonad of Sepiα o 沸 o 乞咒 α だs has been generally be only in vertebrates are apParent − shown to synthesize steroids （Carreau present and Drosdowsky ，1977 ）． The role of sex ly found in highly evolved land snails ． steroid hormones in reproduction is un − − Acknowledgements ’ The author wishes certain ． However ， a negative feedback his Prof ． from the gonad to emission of the optic to express cordial gratitude to gland hormone is suggested by cas 七ration J． Ishida for his eontinuous guidance experiments （Richard and Lamaire ， and encouragement throughout the 1975 ）． course of the present work ． Thanks are also to the ． ． and Reproductive behavior of cephalopods due late Prof Iw Taki − Prof ． T ． Habe Tokai University for is independent of the optic gland hor ， mone ． Even castrated animals show a their inむerest and guidance in this normal sexual behavior． From these reVlew ． findings， the Cephalopoda appears ex ＿ 要 約 − ceptional in their neuroendocrine me ． chanisms ． 本 総 説 で は ，．本邦産 の ミ ．ス ジ マ イ マ イ や コ ウ ラ ナ メ ク ジ の に 牛 It is ge 皿 erally accepted that among な ど 生 殖現象 を 中 心 ，陸 有肺 ホ ル モ ン つ い て し た gastropods the terrestrial forms have 類 の 生殖 の 機構 に 概説 。 生 − ． の reached the highest leveL This is ap 殖現 象 と し て は ，雌 雄 同体 性 ，生 殖 器官 系 分 一 ・ 産 卵 な ど の 連 の 事 象 を parent from their concentrated central 化 成．熟 ，交尾 行動 ， nervous system and complexity of ヒ た 。 ， the 取 リ げ つ い て ホ ル モ ン reproduction ． In the highest group Qf は じ め に ，内 分 泌 系 に ，神 経 a first の 分泌 源 で あ る 脳 の 神 経 分 泌 細 胞 ，内 分泌 器 官 pulmonate gastropods ， and a second order neuroendocrine mechanisms と し て ，脳 の 背方 に 位 置 す る 背 脳 体，視柄 近 く つ い て for the eontrol of male and female ac − に あ る 眼 腺 お よ び 生 殖腺で あ る 両性 腺 に べ で tivities have been de皿 onstrated ． Of 述 た 。 と りわ け，両性腺 は 哺 乳 類 と 同 じ 性 ． − ス テ ロ ドホ ル モ ン が さ 生 期 に 対 great importance for comparative endo イ 産生 れ ， 殖時 て る こ と た こ の crinology is the fact that the more ad − 応 し 挙動す を あげ 。 な お ， れ ら ． ． の よ る の で ホ ル vanced land slugs and snails produce 器 官 の 発達 状態 は 種 に り異 な ， モ ソ 一 で い が る gonadal hormones ， in particular sex 機構 も統 的 に 説明 きな 面 あ 。 一 steroid hormones ， Thus ， reproduction in 次 に ，生 殖 器 官 系 の 般 的 な 形 態 を ，雌 雄 同 ．． some land snails is controlled not only 体性 の 異 な る 両 性 腺，蛋 白 ．腺 お よ び 生 殖輸 管 な こ の の で by a 丘 rst or a second order but also by ど に み る と 共 に ， れ ら 器 官が 発 生 過 程 一 NII-ElectronicN 工 工 Eleotronio Library Service The malaoologioalmalacological societysooiety of Japan ’ 126 VENUS ： Vol ，48 No ．2 （1989．） の の ホ ノレ モ ン の Arch S and 1985 Neuro − 未 分 化 状 態 か ら ， ど よ う な 影 響 ， ． Gainer，H ， ． の か べ secretion In ： A Lajtha ed H α n 〔1− を 受 け て 成 熟す る を 述 た 。 す な わ ち ，神 ． ． り 経 ホ ル モ ソ や 眼 腺 ホ ル モ ン は 両 性 腺 に 刺 激 ホ ル b〔｝ok o／ Neuroch ・emistry ． Vol ．8， PP ． ’ モ f と し て 作 用 し ，前 者 は 雌 性 ，後 者 は 雄 性 の 281 −307，Plenum Publishing Corpora ． ス テ ロ ホ ル モ ソ York 性 イ ド を 産生 さ せ ， 各附 属器 官 tion 、New ． ’ 一 の 成 熟 お よ び 配偶 子 形 成 に 関 5す る 。 ま た ，背 Arnold ，J ．M ，1984 ． Cephalopods ． ∬n ： 脳 ホ ル モ ン が 器 を し て い る 種 A S Tompa N H Verdonk ＆ J A M 体 雌性 附 属 官 制御 ． ． ， ． ． ． ． ． る こ の こ に van Biggelaar もあ 。 れ ら とを比較 内 分泌 学 的 考 察 den eds ，， The Mollusca ． に − し た 。 さ らに ，生 殖行 動 頭 瘤 か ら 放 出 さ れ る Voi ． 7． Reproductioti ． pp ．419 454 ． フ P モ ン ． 性 r ． が 関 ケす る こ と を ，行動 解 発 の 神 Academic Press ， Orlando ． め て べ た Aubry Phermaphro − 経 機構 を 含 述 。 ， R ，1961． Etude de − 系 統発 生上 t 下 位 に 位 置す る 種 で は 牛 殖 現象 ditfsm et de Paction Pharmacodynami の 制御 は 神 経 ホ ル モ ソ に よ る が ，系 統 が E位 に que des hormones de vertebres chez 進 み 体制 が 複雜 化 す る に 伴 い ，上 皮性 の 内分泌 les gasteropodes pulmon6s ． Arch 、 器官 が分 化 し 制 御 の 様 相 に も様 々 な 変 化 が み ∠1？z 〔露 Mic ア 08c MOTphel Exp 50 ： ， ． ． ． ．， る よ こ − ら れ う に な る 。 そ れ ゆ え ， 最後 に ， れ ら 521 602． の こ と を ，軟体 動 物 に お げ る 環境適 応 ，系 統 進 Bailey ， T ． G ． 1973． The 痂 ？フ鷺γ o cul − 化 と 関 連 づ け て 論 じ た 。 ture of reproductive organs in the で ． ・ 木 総 説 は ，単 に 陸 生 貝類 に 留 ま ら ず ，軟体 slug ， A γ iolim α x re 亡iculatus （MUll ）． ノ − 動物 全 般 に 渡 り ．L殖 現 象 お よ び そ の 制御 機 構 の tVeth ．」． Zool 、，23 ： 72 85． 理 解 が 得 ら れ る よ う文 献 を 配 置 し た 。 Barr A R 1928 Some notes on the ， ， ． ． mucous and skin glands of Arion a ．ter． Reference8 ’ ’ ・7 t． ， Mie7 osc ． S （3菰 71 ： 503 −525 Qua 」 ， ． − Abeloos， M ．1943 ． Effects de la castra Beard M ． Millecchia ， C ． ， ， ，L Masuoka ， ・ ・ ． tion chez un mollusque Lim α x m α x i7nus and of ， Arch ．S ．1982 ． Ultrastructure ， L C ．R ． Hebd ． Sea n ，c ． Ac α d ． Sci．，216 ： I ecretion in the atrial gland of a ＿ ・ 90 91 ． mollusc （Aplysia ）． T iss． Cel ら 14 ： E ．1898 ． La fossette − Andre ， triangulaire 297 308． caudal des Arions ． Rei ，．8 〜．｛・isse Zoo 廴 and L 、 1968 − ， Berry ， A ．J ． Chan ， C ． ． Re − 5 ： 179 182 ． productive eondition and tentacle ex − Antheunisse ， L J ．1963． Neurosecretory tirpation in Malayan Aeh α tinα プ祝 伽 α ・− − phenomena in the zebra mussel ，Dre is （Plllmonata ）． Attst．」． Zool．，16 ： 849 ・ sena polV． mot ：）ha， Palls ． A rch ，s Neer 〜． 855 ． ム 16 237 ＿314 − Zoo ， ： ． Bierbauer ， J ．1978 ． Effect of herma Arch S ， Snlock T Gurvis ， ， ， ．， ， R ． and phrodi 七ic gland homogenate on the McCarthy C 1978 ． Atrial − ， gland in regulation of the gametogenesis in of the eg9 − response Helix 蹴 α 擁 α PuI − duction laying in po （Gastropoda ， ・ ・ ’・ Aplysia c α li．for7ticα ．」． Co7np ． Physiol ．， monata ）．・4cta Biol ． Acad ． S 砿 正∬槻 9 ．， − 128 ： 67 70 ， 29 ： 181−187． ． J ． Smock T ． and τld TOr6k L Arch ，S ，Lupatokin ， ， ， Bierbauer， J． a ， ．J， 1968． Beard M 1980 Evidence for an ， ， ． Histopathological examination of the exocrine function of the 4pZ 〃8ぜα atrial optic tentacle with special reference gland ． 」 ， Comp ． Phv． siol ．，141 ： 131− to the morphology of the ．collar and ・ 137 ， lateral cells ． 14c ‘α B iol． Ac α d． Sei． 一 NII-ElectronicN 工 工 Eleotronio Library Service The malacologicalsocietymalacological society of Japan Takecla: Hormonal 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Velige7', 23: 241- Minnen, J. and de Jong-Brink, M. 1976. 244.Chiu, Effeets of castration on the aativity A. Y., Hunkapillar, M. W., Heller, of the endoerine dorsal bodies of the E. Stuart, D.K. Hood, L.E, and freshwater pulmonate snail, Bulinus Strumwasser, F. 1979, Purification tTu?zcatus, intermediate host of Schi- and primary strueture of the neuro- stosto7na haematobium. Neth,, ,J. ZooL, peptide egg-laying hormone of Aplysia 26: 94-105. ealiforniea. PTvc, NatL Aead. Sci., Bonichon, A, 1967. Contribution a 76: 6656-6660. 1'etude de a neuroseeretion et de Choquet, M. 1971. Etude du cycle bio- 1'endoerinologie chez les C6phalopodes. logique et de 1'inversion du sexe chez I, Octop"s vulgaris. Vie et Miliez{, Patelta vulgata L, (Mollusque Gastro- 18: 227-263. pode Prosobranche), Gen. Co･rrep. Bornehen, M. 1967, Untersuehungen Endocrinol., 16: 59-73. zur Sekretion der Fingerformigen Clare, A. S. 1987. Studies on the juxta- 'trochids. DrUsen von Het･ipt pomatia, Z. Zetl- ganglionar organ of ,In: fersch., 78: 402-426. H.H. Boer, W.P.M. Geraerts and J, Bouillon, F. 1956, Infiuence of tem- Joosse eds., Nete'robiotogy, Mollu.scan perature on the histologieal evolution Moclels. North-Holland Publishing of the evetestis of Cepaea nemoTalis. Company, Amsterdam, pp.342-349. Nature, 117; 142-143. Cook, H. 1966, Morphology and his- Bride, J. and Griffond, B. 1981. Etude tology of the central nervous system eomparee de la gonade d'Hetix aspe?'sa of S?tecinea putwis (L), Ao'eh. Neerl. temoins et tentaculectomis6s. Gen, Zool,, 17: 1-72, Comp. Endocrinol., 45: 527-532. de Jong-Brink, M., Schot, L,P.C. Bride, J., Zribi, R, and Gomot, L. 1986. Schoenmaker, H.J,N. and Bergarnin- Action de l'extrait de tentae]es oeu- Sassen, M.J,M. 198L A biochemical laires d'Eseargots juveniles et adultes and quantitative eleetron-mieroseope sur le d6velopment in vivo de la g]ande study on steroidgenesis in ovotestis h albumen d'Helia: aspersa. 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Perga- period on the time schedule of egg- mon Press, Oxford. pp.105-140. mass produetion in Lymnaea stag- Geraerts, W.P.M, 1976, The role of nalis, as induced by ovulation hormone the lateral lobes in the eontrol of injections, Gen. Comp. Endocrinol., growth and reproduction in the her- 49: 255-260. maphrodite freshwater snail, Ly7nnaea Ebberink, R.H.M., van Loenhout, H., stagnalis. Gen. Comp. EndocTinol., Geraerts, W. P, M. and Joosse, J. 1985, 29: 97-108, Purifieatien and amino acid sequenee Geraerts, W. P, M, 1986. Environmental of the ovulation neurohormone ef endoerinology and the eontrol of the Lymnaea stagnalis. Proc. Natl. Acad. reproductive season in molluscs. Adv. Sei., 82: 7767-7771. Inverteb. Rep., 4: 151-162. Emura, S. 1981. Vber Selbstbefrueh- Geraerts, W. P. M, and Joosse, J. 1975, tung Lungenschneeken. bei Butl. Ni･i- Control of vitellogenesis and growth gata Seiryo Woman Coll., 11: 17-36 of female aeeessory sex organs by the (In Japanese). dorsal body horrnone (DBH) in the Feral, C., le Gall, S,. Marie-Christine, hermaphrodite freshwater snail, M. and Lengronne, C. 1987. The Lymnaea sta･gnalis. Gen. Comp. Endo- neuroendocrine mechanism responsible crinel., 27: 450-464. for sexual inversion of the gonad in Geraerts, W. P. M. and Bohlken, S. 1976, the protandrous hermaphrodie mol- The eontrol of ovulation in the her- luscs, Crepidula fornicata. L, Gen. maphroditic freshwater snail, Lymnaea Comp. Endocrinol., 65: 432-438. stagnalis by the neurohormone of the Filhol, J, 1938. Reeherches sur la na- caudodersal cells. Gen. Comp. Endo- ture des lepidosornes et les ph6nom6nes c･rinol., 28: 350-357, eytologiques de la secretion chez les Geraerts, W.P.M, and Joosse, J. 1984. gastropodes pulmones. III. Vesieles Freshwater snail, In: A,S. Tompa, mu]tifides et canal d6ferent. Arch. N. H. Verdonk and J. 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Bioehe?n. a. Steroids of inverbrates IV. On PhysioL, 84A: 289-294. the optic tentacle-gonadal axis in the Haskins, J. T., Price, C. H. and Blanken- eontrol of the male-phase ovotestis in ship, J. E, 1981. A light and eleetron the slug (Ariolimax cali.fornicus). microseopie investigation of the neuro- Gen. Comp. Endocn'nol., 15: 101-119, secretory bag eells of Aplysin. J. Gottfried, H, and Dorfman, R.J, 1970b. Neurocytol., 10: 729-747. Steroids of invertebrates V. The in Herbert, D.G. 1982. Fine structural vit･ro biosynthesis of steroids by the observations on the juxtaganglionar NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 130 VENUS: VoL 48,No, 2 (1989) organ of Gibbula u?nbilicalis. Moll, man & M. Porchet eds., Biosynthesis, Stud,, 226-228, Metabolism and Mode ef Action of Heller. E., Kaezmarek, K., Hunkapillar, Invertebrate Hormones, pp. 149-172, M.W. Hood, L.E, and Strumwasser, Springer-Verlag, Berlin. F. 1980. Purification and primary Joosse,J. 1987. Neuroendoerine integra- strueture of two neuroactive peptides tion: Molluscan models. Proc. Jap. that eause bag cell afterdischarge and Soc. Comp. Endocrinol., 2: 7-10. egg-laying in Aptys･ia. Proc. Natl. Jeosse, J. 1988. The hormones of mol- Acaal. Sci., 77: 2328-2332, luscs. In: H. Laufer & G.H, Downer Herlin-Houtteville, P. and Lubet, P.E. eds., Invertebrate Endocrinology. Vol. 1975. The sexuality of pelecypod 2. Endocrinology of Selected htver- molluscs, ht: R. Reinboth ed., Inter- tebrate Types. pp.89-140. Alan R. sexuathtll in the Aniwaal K･ingdom. Liss, New York. pp,179-187, Springer-Verlag, Berlin, Joosse, J. and Geraerts, W.P. M, 1983, Herzberg, F. and Herzberg, A. 1962. Endoerinology. In: A.S.M. Saleud- Observations on reproduction in "elix dine & KM. Wilbur, eds., The Mot- aspeTsa. Am, MidL Nat., 68: 297-306. lusca. Vol. 4. Physiotogy. Part 1. pp. Hughes, G.M. and Kerkut, G.A, 1956. 317-406, Academic Press, Orleando. Electrieal aetivity in a slug ganglion Joesse, J,, Ebberink, R. H,M., Geraerts, in relation to the concentration of W. P. M., Jansen. R. F. and Maat, A. Loeke solution. .J. Exp. Biol., 33: ter. 1985. The eaudo-dorsal eells of 282L294. bymnaea stagnalis: Primary strueture Ikeda, K. 1937. Cytogenetie studies en of the ovulation hormone and biosyn- the self-fertilization of Philomyctts thesis and role of multiple peptides bilineatus Benson. J. Sci, Hiroshi"za released during egg-laying. In: H, Univ. Se･r. B. Div, 1,, 5: 67-123, Kobayashi, H.A. Bern & A. Urano Jeppesen, L. 1976. The control of mat- eds., Nezarosecretion azad Biology o.t' ing behayior in Hetix pomat･ia. Anim Neuropeptides, pp.50-59, Jap. Sci, Behav., 24: 275-290. Soc. Press, Tokyo. Joosse, J. 1975. 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