Gonadotropin-releasing hormone (GnRH) is widely distributed in the brain of fishes where it may function as a neuromodulator of sensory processing and behavior. Immunocytochemical and neuronal label experiments were conducted on species from four families of coral reef fishes (Chaetodontidae, butterflyfish; Pomacentridae, damselfish; Gobiidae, goby; and Labridae, wrasse) to assess conservation of GnRH targets in the visual processing retina and brain. In all species, GnRH-immunoreactive (-ir) axons from the terminal nerve project principally to the boundary between the inner plexiform (IPL) and inner nuclear (INL) layers of the retina, and are less prominent in the optic nerve, ganglion cell, IPL and INL. However, the density of GnRH innervation within the retina differed among fish species with highest concentrations in the damselfish and butterflyfish and lowest in the goby and wrasse. Experiments also show that GnRH receptors are associated with GnRH-ir axons within the fish retina primarily at the IPL-INL boundary, the region of light-dark adaptation and image processing of contrast, motion or color. GnRH-ir axons overlapped central projections of retinal ganglion cell axons primarily within the stratum album centrale and stratum griseum centrale of the tectum in all species, and were concentrated in several diencephalic visual processing centers. GnRH receptors are also localized to diencephalic visual centers and the stratum griseum periventriculare of the tectum, where motion perception and coordination of motor behavioral responses in three-dimensional space occur. This work demonstrates that the basic neural substrates for peptide-sensory convergence are conserved at multiple processing levels in the visual system of several reef fishes. Species differences in GnRH innervation to the retina and GnRH receptor distributions may be related to phylogeny, their use of vision in natural behaviors, or possibly binding properties of the antibodies. Future studies are needed to characterize the exact GnRH variants and receptor types found in these species so that possible functional consequences of GnRH influence on vision can be defined.

1.
Abe H, Oka Y (2002) Mechanisms of the modulation of pacemaker activity by GnRH peptides in the terminal nerve-GnRH neurons. Zool Sci 19:111–128.
2.
Amano M, Urano A, Aida K (1997) Distribution and function of gonadotropin-releasing hormone (GnRH) in the teleost brain. Zool Record 14:1–11.
3.
Ball AK, Stell WK, Tutton DA (1989) Efferent projections to the goldfish retina. NATO ASI Series, H31:103–116.
4.
Barchas JD, Akil H, Elliott GR, Holman RB, Watson SJ (1978) Behavioral neurochemistry: neuroregulators and behavioral states. Science 200:964–973.
5.
Barry KL, Hawryshyn CW (1999) Effects of incident light and background conditions on potential conspicuousness of Hawaiian coral reef fish. J Mar Biol Assoc UK 79:495–508.
6.
Bass AH, Grober MS (2001) Social and neural modulation of sexual plasticity in teleost fish. Brain Behav Evol 5:293–300.
7.
Behrens UD, Wagner HJ (2004) Terminal nerve and vision. Micro Res Tech 65:25–32.
8.
Behrens UD, Douglas RH, Wagner HJ (1993) Gonadotropin-releasing hormone, a neuropeptide of efferent projections to the teleost retina induces light-adaptive spinule formation on horizontal cell dendrites in dark-adapted preparations kept in vitro. Neurosci Lett 164:59–62.
9.
Bennett AT, Cuthill IC, Partridge JC, Lunau K (1997) Ultraviolet plumage colors predict mate preferences in starlings. Proc Natl Acad Sci USA 94:8618–8621.
10.
Braford Jr MR, Northcutt RG (1983) Organization of the diencephalon and pretectum of the ray finned fishes. In: Fish Neurobiology (Davis RE, Northcutt RG, eds), pp 117–164. Ann Arbor, MI.: University of Michigan.
11.
Butler AB, Hodos W (1996) Comparative Vertebrate Neuroanatomy: Evolution and Adaptation. Wiley-Liss, Inc., New York, NY.
12.
Chartrel N, Collin F, Huang Y, Montero M, Tonon M, Goos HJT, Dufour S, Vaudry H (1998) Characterization and localization of two forms of gonadotropin-releasing hormone (GnRH) in the spinal cord of the frog Rana ridibunda. Cell Tiss Res 293:235–243.
13.
Chen C, Fernald RD (2006) Distributions of two gonadotropin-releasing hormone receptor types in a cichlid fish suggest functional specialization. J Comp Neurol 495:314–323.
14.
Eisthen HL, Delay RJ, Wirsig-Wiechmann CR, Dionne VE (2000) Neuromodulatory effects of gonadotropin releasing hormone on olfactory receptor neurons. J Neurosci 20:3947–3955.
15.
Ekstrom P (1984) Central neural connections of the pineal organ and retina in the teleost Gasterosteus aculeatus L. J Comp Neurol 226:321–335.
16.
Elofsson UOE, Winberg S, Francis RC (1997) Number of preoptic GnRH-immunoreactive cells correlates with sexual phase in a protandrously hermaphroditic fish, the dusky anemonefish (Amphiprion melanopus). J Comp Physiol A 181:484–492.
17.
Elofsson UOE, Winberg S, Nilsson GE (1999) Relationships between sex and the size and number of forebrain gonadotropin-releasing hormone-immunoreactive neurons in the Ballan wrasse (Labrus berggylta), a protogynous hermaphrodite. J Comp Neurol 410:158–170.
18.
Fernald RD, White RB (1999) Gonadotropin-releasing hormone genes: phylogeny, structure, and functions. Front Neuroendocrinol 20:224–240.
19.
Foran CM, Myers DA, Bass AH (1997) Modification of gonadotropin releasing hormone (GnRH) mRNA expression in the retinal-recipient thalamus. Gen Comp Endocrinol 106:251–264.
20.
Forlano PM, Maruska KP, Sower SA, King JA, Tricas TC (2000) Differential distribution of gonadotropin-releasing hormone-immunoreactive neurons in the stingray brain: functional and evolutionary considerations. Gen Comp Endocrinol 118:226–248.
21.
Fukuda FM, Ishimoto I, Shiosaka S, Kuwayama Y, Shimizu Y, Inagaki S, Sakanaka M, Takagi H, Senba E, Takatsuki K, Umegaki K, Tohyama M (1982) Localization of LH-RH immunoreactivity in the avian retina. Curr Eye Res 2:71–74.
22.
Fuxe K, Agnati LF (1991) Two principal modes of electrochemical communication in the brain: volume versus wiring transmission. In: Volume Transmission in the Brain: Novel Mechanisms for Neural Transmission (Fuxe K, Agnati LF, eds), pp 1–9. New York: Raven Press.
23.
Green CB, Besharse JC (2004) Retinal circadian clocks and control of retinal physiology. J Biol Rhythm 19:91–102.
24.
Grens KE, Greenwood AK, Fernald RD (2005) Two visual processing pathways are targeted by gonadotropin-releasing hormone in the retina. Brain Behav Evol 66:1–9.
25.
Grober MS (1998) Socially controlled sex change: integrating ultimate and proximate levels of analysis. Acta Ethol 1:3–17.
26.
Grober MS, Bass AH (1991) Neuronal correlates of sex/role change in labrid fishes: LHRH-like immunoreactivity. Brain Behav Evol 38:302–312.
27.
Grober MS, Bass AH, Burd G, Marchaterre MA, Segil N, Scholz K, Hodgson T (1987) The nervus terminalis ganglion in Anguilla rostrata: an immunocytochemical and HRP histochemical analysis. Brain Res 436:148–152.
28.
Guilgur LG, Moncaut NP, Canario AVM, Somoza GM (2006) Evolution of GnRH ligands and receptors in gnathostomata. Comp Biochem Physiol A 144:272–283.
29.
Haneda K, Oka Y (2004) Selective modulation of voltage-gated calcium channels in the terminal nerve gonadotropin-releasing hormone neurons of a teleost, the dwarf gourami (Colisa lalia). Endocrinology 145:4489–4499.
30.
Helfrich P (1958) The early life history and reproductive behavior of the maomao, Abudefduf abdominalis (Quoy and Gaimard). Ph.D. Dissertation, Zoology Dept., University of Hawaii at Manoa, 228pp.
31.
Hourigan TF (1987) The behavioral ecology of three species of butterflyfishes. Ph.D. Dissertation, Zoology Dept., University of Hawaii at Manoa. 496 pp.
32.
Illing N, Troskie BE, Nahorniak CS, Hapgood JP, Peter RE, Millar RP (1999) Two gonadotropin-releasing hormone receptor subtypes with distinct ligand selectivity and differential distribution in brain and pituitary in the goldfish (Carassius auratus). Proc Natl Acad Sci USA 96:2526–2531.
33.
Jan YN, Jan LY (1983) A LHRH-like peptidergic neurotransmitter capable of ‘action at a distance’ in autonomic ganglia. Trends Neurosci 6:320–325.
34.
Jennes L, Eyigor O, Janovick JA, Conn PM (1997) Brain gonadotropin releasing hormone receptors: localization and regulation. Recent Prog Horm Res 52:475–490.
35.
Jennes L, Conn PM (1993) Gonadotropin-releasing hormone and its receptors in rat brain. Front Neuroendocrinol 15:51–77.
36.
Jennes L, Dalati B, Conn PM (1988) Distribution of gonadotropin releasing hormone agonist binding sites in the rat central nervous system. Brain Res 452:156–164.
37.
Jodo A, Ando H, Urano A (2003) Five different types of putative GnRH receptor gene are expressed in the brain of Masu salmon (Oncorhynchus masou). Zool Sci 20:1117–1125.
38.
Kodric-Brown A, Johnson SC (2002) Ultraviolet reflectance patterns of male guppies enhance their attractiveness to females. Anim Behav 63:391–396.
39.
LeBas NR, Marshall NJ (2000) The role of colour in signaling and male choice in the agamid lizard Ctenophorus ornatus. Proc R Soc Lond B 267:445–452.
40.
Lethimonier C, Madigou T, Munoz-Cueto J, Lareyre J, Kah O (2004) Evolutionary aspects of GnRHs, GnRH neuronal systems and GnRH receptors in teleost fish. Gen Comp Endocrinol 135:1–16.
41.
Losey GS, McFarland WN, Loew ER, Zamzow JP, Nelson PA, Marshall NJ (2003) Visual biology of Hawaiian coral reef fishes. I. ocular transmission and visual pigments. Copeia 2003:433–454.
42.
Losey GS, Cronin TW, Goldsmith TH, Hyde D, Marshall NJ, McFarland WN (1999) The UV world of fishes: a review. J Fish Biol 54:921–943.
43.
Maaswinkel H, Li L (2003) Olfactory input increases visual sensitivity in zebrafish: a possible function for the terminal nerve and dopaminergic interplexiform cells. J Exp Biol 206:2201–2209.
44.
Madigou T, Mananos-Sanchez E, Hulshof S, Anglade I, Zanuy S, Kah O (2000) Cloning, tissue distribution, and central expression of the gonadotropin-releasing hormone receptor in the rainbow trout (Oncorhynchus mykiss). Biol Reprod 63:1857–1866.
45.
Medina M, Reperant J, Miceli D, Ward R, Arckens L (2005) GnRH-immunoreactive centrifugal visual fibers in the Nile crocodile (Crocodylus niloticus). Brain Res 1052:112–117.
46.
Meredith M, Fernandez-Fewell G (1994) Vomeronasal system, LHRH, and sex behavior. Psychoneuroendocrinology 19:657–672.
47.
Moncaut N, Somoza GM, Power D, Canario A (2005) Five gonadotropin-releasing hormone receptors in a teleost fish: isolation, tissue distribution and phylogenetic relationships. J Mol Endocrinol 34:767–779.
48.
Moore FL, Rose JD (2002) Sensorimotor processing model: how vasotocin and corticosterone interact and control reproductive behavior in an amphibian. In: Hormones, Brain and Behavior. Vol. II (Pfaff D, Arnold A, Etgen A, Fahrbach S, Rubin R, eds), pp 515–544. New York: Elsevier Science.
49.
Munz H, Class B, Stumpf WE, Jennes L (1982) Centrifugal innervation of the retina by luteinizing hormone releasing hormone (LHRH) – immunoreactive telencephalic neurons in teleostean fishes. Cell Tiss Res 222:313–323.
50.
Munz H, Stumpf WE, Jennes L (1981) LHRH systems in the brain of platyfish. Brain Res 221:1–13.
51.
Muske LE (1993) Evolution of gonadotropin-releasing hormone (GnRH) neuronal systems. Brain Behav Evol 42:215–230.
52.
Muske LE, Fernald RD (1987) Control of a teleost social signal. I. Neural basis for differential expression of a color pattern. J Comp Physiol A 160:89–97.
53.
Myrberg AA, Fuiman LA (2002) The Sensory World of Coral Reef Fishes. In: Coral Reef Fishes (Sale P, ed), pp 123–148. New York:Elsevier Science.
54.
Myrberg AA, Thresher RE (1974) Interspecific aggression and its relevance to the concept of territoriality in reef fishes. Am Zool 14:81–96.
55.
Northcutt RG (1983) Evolution of the optic tectum in ray-finned fishes. In: Fish Neurobiology (Davis RE, Northcutt RG, eds), pp 1–42. Ann Arbor, MI.: University of Michigan Press.
56.
Oka Y (1997) GnRH neuronal system of fish brain as a model system for the study of peptidergic neuromodulation. In: GnRH Neurons: Gene to Behavior (Parhar IS, Sakuma Y, eds), pp 245–276. Brain Shuppan: Tokyo, Japan.
57.
Oka Y (1992) Gonadotropin-releasing hormone (GnRH) cells of the terminal nerve as a model neuromodulator system. Neurosci Lett 142:119–122.
58.
Oka Y, Matsushima T (1993) Gonadotropin-releasing hormone (GnRH)-immunoreactive terminal nerve cells have intrinsic rhythmicity and project widely in the brain. J Neurosci 13:2161–2176.
59.
Oka Y, Ichikawa M (1992) Ultrastructural characterization of gonadotropin-releasing hormone (GnRH)-immunoreactive terminal nerve cells in the dwarf gourami. Neurosci Lett 140:200–202.
60.
Oka Y, Ichikawa M (1990) Gonadotropin-releasing hormone (GnRH) immunoreactive system in the brain of the dwarf gourami (Colisa lalia) as revealed by light microscopic immunocytochemistry using a monoclonal antibody to common amino acid sequence of GnRH. J Comp Neurol 300:511–522.
61.
Okubo K, Suetake H, Usami T, Aida K (2000) Molecular cloning and tissue-specific expression of a gonadotropin-releasing hormone receptor in the Japanese eel. Gen Comp Endocrinol 119:181–192.
62.
Parhar IS, Soga T, Sakuma Y, Millar RP (2002) Spatio-temporal expression of gonadotropin-releasing hormone receptor subtypes in gonadotropes, somatotropes and lactotropes in the cichlid fish. J Neuroendocrinol 14:657–665.
63.
Park D, Eisthen HL (2003) Gonadotropin-releasing hormone (GnRH) modulates odorant responses in the peripheral olfactory system of Axolotls. J Neurophysiol 90:731–738.
64.
Peter RE, Prasada Rao PD, Baby SM, Illing N, Millar RP (2003) Differential brain distribution of gonadotropin-releasing hormone receptors in the goldfish. Gen Comp Endocrinol 132:399–408.
65.
Presson J, Fernald RD, Max M (1985) The organization of retinal projections to the diencephalon and pretectum in the cichlid fish, Haplochromis burtoni. J Comp Neurol 235:360–374.
66.
Privitera LA (2002) The reproductive behavior and ecology of a coral-reef gobiid fish, Asterropteryx semipunctata. PhD. Dissertation, Zoology Dept., University of Hawaii at Manoa, 144 pp.
67.
Reese ES (1975) A comparative field study of the social behavior and related ecology of reef fishes of the family Chaetodontidae. Z Tierpsychol 37:37–61.
68.
Reperant J, Ward R, Miceli D, Rio JP, Medina M, Kenigfest NB, Vesselkin NP (2006) The centrifugal visual system of vertebrates: a comparative analysis of its functional anatomical organization. Brain Res Rev 52:1–57.
69.
Robison RR, White RB, Illing N, Troskie BE, Morley M, Millar RP, Fernald RD (2001) Gonadotropin-releasing hormone receptor in the teleost Haplochromis burtoni: structure, location, and function. Endocrinology 142:1737–1743.
70.
Rose JD, Moore FL (2002) Behavioral neuroendocrinology of vasotocin and vasopressin and the sensorimotor processing hypothesis. Front Neuroendocrinol 23:317–341.
71.
Rosen G, Sherwood N, King J (1997) Immunoreactive gonadotropin-releasing hormone (GnRHir) is associated with vestibular structures in the green anole (Anolis carolinensis). Brain Behav Evol 50:129–138.
72.
Ross RM (1982) Sex change in the endemic Hawaiian labrid Thalassoma duperrey (Quoy and Gaimard): a behavioral and ecological analysis. Ph.D. Dissertation, Zoology Dept., University of Hawaii at Manoa, 171 pp.
73.
Rowland WJ, Grindle N, Maclaren RD, Granquist R (2002) Male preference for a subtle posture cue that signals spawning readiness in female sticklebacks. Anim Behav 63:743–748.
74.
Sakuma Y, Suga S (1997) GnRH and female rat sexual behavior. In: GnRH Neurons: Gene to Behavior (Parhar IS, Sakuma Y, eds), pp 389–400. Tokyo, Japan: Brain Shuppan.
75.
Soga T, Ogawa S, Millar RP, Sakuma Y, Parhar IS (2005) Localization of the three GnRH types and GnRH receptors in the brain of a cichlid fish: insights into their neuroendocrine and neuromodulator functions. J Comp Neurol 487:28–41.
76.
Springer AD (1983) Centrifugal innervation of goldfish retina from ganglion cells of the nervus terminalis. J Comp Neurol 214:404–415.
77.
Stell WK, Walker SE, Ball AK (1987) Functional-anatomical studies on the terminal nerve projection to the retina of bony fishes. Ann NY Acad Sci 80–96.
78.
Stell WK, Walker SE, Chohan KS, Ball AK (1984) The goldfish nervus terminalis: a luteinizing hormone-releasing hormone and molluscan cardioexcitatory peptide immunoreactive olfactoretinal pathway. Proc Natl Acad Sci USA 81:940–944.
79.
Tricas TC (1989) Food and competitors as determinants of territory size in the Hawaii butterflyfish, Chaetodon multicinctus. Anim Behav 37:830–841.
80.
Umino O, Dowling JE (1991) Dopamine release from interplexiform cells in the retina: effects of GnRH, FMRFamide, bicuculline, and enkephalin on horizontal cell activity. J Neurosci 11:3034–3046.
81.
Walker SE, Stell WK (1986) Gonadotropin-releasing hormone (GnRH), molluscan cardioexcitatory peptide (FMRFamide), enkephalin and related neuropeptides affect goldfish retinal ganglion cell activity. Brain Res 384:262–273.
82.
Wirsig-Wiechmann CR (1993) Peripheral projections of nervus terminalis LHRH-containing neurons in the tiger salamander, Ambystoma tigrinum. Cell Tiss Res 273:31–40.
83.
Wirsig-Wiechmann CR (2001) Function of gonadotropin-releasing hormone in olfaction. Keio J Med 50:81–85.
84.
Wirsig-Wiechmann CR, Lepri JJ (1991) LHRH-immunoreactive neurons in the pterygopalatine ganglia of voles: a component of the nervus terminalis? Brain Res 568:289–293.
85.
Wirsig-Wiechmann CR, Oka Y (2002) The terminal nerve ganglion cells project to the olfactory mucosa in the dwarf gourami. Neurosci Res 44:337–341.
86.
Wirsig-Wiechmann CR, Wiechmann AF (2002) Vole retina is a target for gonadotropin-releasing hormone. Brain Res 950:210–217.
87.
Zucker CL, Dowling JE (1987) Centrifugal fibres synapse on dopaminergic interplexiform cells in the teleost retina. Nature 330:166–168.
88.
Zumpe D (1965) Laboratory observations on the aggressive behavior of some butterflyfishes (Chaetodontidae). Z Flyerpsychol 22:226–244.
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