Professor Clive Coen - Publications: Kalamatianos, T., Faulkes, C.G., Oosthuizen, M.K., Poorun, R., Bennett, N.C. & Coen, C.W. (2010). Telencephalic binding sites for oxytocin and social organization: a comparative study of eusocial naked mole-rats and solitary cape mole-rats. Journal of Comparative Neurology 518, 1792-1813. Vida, B., Deli, L., Hrabovszky, E., Kalamatianos, T., Caraty, A., Coen, C.W., Liposits, Z. & Kalló, I. (2010). Evidence for suprachiasmatic vasopressin neurones innervating kisspeptin neurones in the rostral periventricular area of the mouse brain: regulation by oestrogen. Journal of Neuroendocrinology 22, 1032-1039. Samuelsson, A.M., Morris, A., Igosheva, N., Kirk, S.L., Pombo, J.M., Coen, C.W., Poston, L. & Taylor, P.D. (2010). Evidence for sympathetic origins of hypertension in juvenile offspring of obese rats. Hypertension 55, 76-82. Kirk, S.L., Samuelsson, A.M., Argenton, M., Dhonye, H., Kalamatianos, T., Poston, L., Taylor, P.D. & Coen, C.W. (2009). Maternal obesity induced by diet in rats permanently influences central processes regulating food intake in offspring. Public Library of Science (PLoS) One 4, e5870 (1-13). Kalamatianos, T., Grimshaw, S.E., Poorun, R., Hahn, J.D. & Coen, C.W. (2008). Fasting reduces KiSS-1 expression in the anteroventral periventricular nucleus (AVPV): effects of fasting on the expression of KiSS-1 and NPY in the AVPV and/or arcuate nucleus of female rats. Journal of Neuroendocrinology 20, 1089-1097. Oosthuizen, M.K., Bennett, N.C., Lutermann, H. & Coen, C.W. (2008). Reproductive suppression and the seasonality of reproduction in the social Natal mole-rat (Cryptomys hottentotus natalensis). General and Comparative Endocrinology 159, 236-40. Oosthuizen, M.K., Bennett, N.C. & Coen, C.W. (2008). An immunohistochemical study of the gonadotrophin-releasing hormone 1 system in solitary Cape mole-rats, Georychus capensis, and social Natal mole-rats, Cryptomys hottentotus natalensis. Neuroscience 157, 164-173. Vida, B., Hrabovszky, E., Kalamatianos, T., Coen, C.W., Liposits, Z. & Kalló, I. (2008). Oestrogen receptor α and β in the suprachiasmatic nucleus of male and female mice. Journal of Neuroendocrinology 20, 1270-1277. Dakin, C.L., Wilson, C.A., Kalló, I., Coen, C.W. & Davies, D.C. (2008). Neonatal stimulation of 5-HT2 receptors reduces androgen receptor expression in the rat anteroventral periventricular nucleus and sexually dimorphic preoptic area. European Journal of Neuroscience 27, 2473-2480. Hart, L., Bennett, N.C., Kalamatianos, T., Oosthuizen M.K., Jarvis, J.U.M., O’Riain, M.J. & Coen, C.W. (2008). Neuroanatomical investigation of the gonadotrophin-releasing hormone 1 system in the seasonally breeding Cape dune mole-rat, Bathyergus suillus. Brain Research Bulletin 77, 185-188. Oosthuizen, M.K., Lutermann, H., Bennett, N.C. & Coen C.W. (2008). Reproductive suppression and the seasonality of reproduction in the social Natal mole-rat (Cryptomys hottentotus natalensis). General and Comparative Endocrinology 159, 236-240. Oosthuizen, M.K., Bennett, N.C. & Coen C.W. (2008). An immunohistochemical study of the gonadotrophin-releasing hormone 1 system in solitary Cape mole-rats, Georychus capensis, and social Natal mole-rats, Cryptomys hottentotus natalensis. Neuroscience 157, 164-173.2 Hahn, J & Coen, C.W. (2006). A comparative study of the sources of neuronal projections to the site of gonadotrophin-releasing hormone perikarya and to the anteroventral periventricular nucleus in female rats. Journal of Comparative Neurology 494, 190-214. Du Toit, L., Bennett, N.C., Katz, A.A., Kalló, I. & Coen, C.W. (2006). Relations between social status and the gonadotrophin-releasing hormone system in females of two cooperatively breeding species of African mole-rats, cryptomys hottentotus hottentotus and cryptomys hottentotus pretoriae: neuroanatomical and neuroendocrinological studies. Journal of Comparative Neurology 494, 303-313. Joy, S., Siow, R.C., Rowlands, D.J., Becker, M., Wyatt, A.W., Aaronson, P.I., Coen, C.W., Jacob, R., Kalló, I., Mann, G.E. (2006). The isoflavone equol mediates rapid vascular relaxation: Ca2+-independent activation of eNOS/Hsp90 involving ERK1/2 and Akt phosphorylation in human endothelial cells. Journal of Biological Chemistry 281, 27335–27345. Murray, J.F., Hahn, J.D., Kennedy, A.R., Small, C.J., Bloom, S.R., Haskell-Luevano, C., Coen, C.W. & Wilson, C.A. (2006). Evidence for a stimulatory action of melanin-concentrating hormone on luteinising hormone release involving MCH1 and melanocortin-5 receptors. Journal of Neuroendocrinology 18, 157–167. Du Toit, L., Bennett, N.C., Gutjahr, G.H. & Coen, C.W. (2006). Reproductive suppression in subordinate female highveld mole-rats (cryptomys hottentotus pretoriae): no role for endogenous opioid peptides. Physiology and Behavior 87, 897-902. Kalamatianos, T., du Toit, L., Hrabovszky, E., Kalló, I., Marsh, P., Bennett, N.C. & Coen, C.W. (2005). Characterization of gonadotrophin releasing hormone precursor cDNA in the Old World mole-rat cryptomys hottentotus pretoriae: high degree of identity with the New World guinea pig sequence. Journal of Neuroendocrinology 17, 265-268. Bhogal, R.K., Mitchell, A.L. & Coen, C.W. (2004). Identification of putative rat ribonuclease III by differential display: a novel rat mRNA expressed in a circadian manner in the rat suprachiasmatic nucleus. Molecular Brain Research 131, 51-57. Threlfell, S., Cragg, S.J., Kalló, I., Turi, G., Coen, C.W. & Greenfield, S.A. (2004). Histamine H3 receptors inhibit serotonin (5-HT) release in substantia nigra pars reticulata. Journal of Neuroscience 24, 8704-8710. Kalló, I., Kalamatianos, T., Piggins, H.D. & Coen, C.W. (2004). Ageing and the diurnal expression of the mRNAs for vasoactive intestinal polypeptide and for the VPAC2 and PAC1 receptors in the suprachiasmatic nucleus of male rats. Journal of Neuroendocrinology 16, 758766. Kalamatianos, T., Kalló, I., Piggins, H.D. & Coen, C.W. (2004). Expression of VIP and PACAP receptor mRNA in peptide synthesising cells within the suprachiasmatic nucleus of the rat and in its efferent target sites. Journal of Comparative Neurology 475, 19-35. Li, X.F., Edward, J., Mitchell, J.C., Shao, B., Bowe, J.E., Coen, C.W., Lightman S.L. & O'Byrne, K.T. (2004). Differential effects of repeated restraint stress on pulsatile luteinizing hormone secretion in female Fischer, Lewis and Wistar rats. Journal of Neuroendocrinology 16, 620-627. Hrabovszky, E., Kalló, I., Steinhauser, A., Merchenthaler, I., Coen, C.W., Petersen, S.L. and Liposits, Zs. (2004). Estrogen receptor-β in oxytocin and vasopressin neurons of the rat and human hypothalami: immunocytochemical and in situ hybridization studies. Journal of Comparative Neurology 473, 315-333.3 Kalamatianos, T., Kalló, I., Goubillon, M.L. & Coen, C.W. (2004). Cellular expression of V1a vasopressin receptor mRNA in the female rat preoptic area: effects of oestrogen. Journal of Neuroendocrinology 16, 525-533. Kalamatianos, T., Kalló, I. & Coen, C.W. (2004). Ageing and the diurnal expression of the mRNAs for vasopressin and for the V1a and V1b vasopressin receptors in the suprachiasmatic nucleus of male rats. Journal of Neuroendocrinology 16, 493-501. Kalló, I., Kalamatianos, T., Wiltshire, N., Shen, S., Sheward, W.J., Harmar, A.J. & Coen, C.W. (2004). Transgenic approach reveals expression of the VPAC2 receptor in phenotypically defined neurons in the mouse suprachiasmatic nucleus and in its efferent target sites. European Journal of Neuroscience 19, 2201-2211. Molteno, A.J., Kalló, I., Bennett, N.C., King, J.A. & Coen C.W. (2004). A neuroanatomical and neuroendocrinological study into the relation between social status and the GnRH system in cooperatively breeding female Damaraland mole-rats Cryptomys damarensis. Reproduction 127, 13-22. Forsling, M.L., Kalló, I., Hartley, D.E., Heinze, L., Ladek, R., Coen, C.W. & File, S.E. (2003). Oestrogen-β receptor and neurohypophysial hormones: functional interaction and neuroanatomical localisation. Pharmacology Biochemistry and Behaviour 76, 535-542. Li, X.F., Mitchell, J.C., Wood, S., Coen, C.W., Lightman, S.L. & O’Byrne, K.T. (2003). The effect of oestradiol and progesterone on hypoglycaemic stress-induced suppression of pulsatile luteinizing hormone release and on corticotrophin-releasing hormone mRNA expression in the rat. Journal of Neuroendocrinology 15, 468-476. Small, C.J., Goubillon, M.L., Murray, J.F., Siddiqui, A., Grimshaw, S.E., Young, H., Sivanesan, V., Kalamatianos, T., Kennedy, A.R., Coen, C.W., Bloom S.R., & Wilson, C.A. (2003). Central orexin A has site-specific effects on luteinizing hormone release in female rats. Endocrinology 144, 3225-3236. Hahn, J.D., Kalamatianos, T. & Coen C.W. (2003). Studies on the neuroanatomical basis for stress-induced oestrogen-potentiated suppression of reproductive function: evidence against direct corticotrophin releasing hormone projections to the vicinity of luteinising hormone releasing hormone cell bodies in female rats. Journal of Neuroendocrinology 15, 732-742. Reed, H.E., Cutler, D.J., Brown, T.M., Brown, J., Coen, C.W. & Piggins, H.D. (2002). Effects of vasoactive intestinal polypeptide on neurones of the rat suprachiasmatic nuclei in vitro. Journal of Neuroendocrinology 14, 639-646. Kalló, I., Butler, J.A., Barkovics-Kalló, M., Goubillon, M.-L. & Coen, C.W. (2001). Oestrogen receptor β-immunoreactivity in gonadotrophin-releasing hormone-expressing neurones: regulation by oestrogen. Journal of Neuroendocrinology 13, 741-748. McGarvey, C., Cates, P.A., Brooks, A., Swanson, I.A., Milligan, S.R., Coen, C.W. & O'Byrne, K.T. (2001). Phytoestrogens and gonadotropin-releasing hormone pulse generator activity and pituitary luteinizing hormone release in the rat. Endocrinology 142, 1202-1208. Reed, H.E., Meyer-Spasche, A., Cutler, D.J., Coen, C.W. & Piggins, H.D. (2001). Vasoactive intestinal polypeptide (VIP) phase-shifts the rat suprachiasmatic nucleus clock in vitro. European Journal of Neuroscience 13, 839-843. Shen, S., Spratt, C., Sheward, W.J., Kalló, I, West, K., Morrison, C.F., Coen, C.W., Marston, H.M. & Harmar, A.J. (2000). Overexpression of the human VPAC2 receptor in 4 the suprachiasmatic nucleus alters the circadian phenotype of mice. Proceedings of the National Academy of Sciences 97, 11575-11580. Goubillon, M.-L., Strutton, P.H., O’Byrne, K.T., Thalabard, J.-C. & Coen, C.W. (1999). Ketamine-induced general anesthesia is compatible with gonadotrophin releasing-hormone pulse generator activity in gonadectomized rats: prospects for detailed electrophysiological studies in vivo. Brain Research 841, 197-201. Fekete, C., Strutton, P.H., Cagampang, F.R.A., Hrabovszky, E., Kalló, I., Shughrue, P.J., Dobó, E., Mihály, E., Baranyi, L., Okada, H., Panula, P., Merchenthaler, I., Coen, C.W. & Liposits, Z. (1999). Estrogen receptors are abundantly expressed in central histaminergic neurons: evidence for a new neuroendocrine pathway regulating luteinizing hormone releasing hormone in rats and humans. Endocrinology 140, 4335-4341. Cagampang, F.R.A., Whatley, S., Mitchell, A.L., Powell, J.F., Campbell, I.C. & Coen, C.W. (1999). Circadian regulation of prion protein mRNA in the rat forebrain: a widespread and synchronous rhythm. Neuroscience 91, 1201-1204. Cagampang, F.R.A., Strutton, P.H., Goubillon, M.-L. & Coen, C.W. (1999). Adrenomedullectomy prevents the suppression of pulsatile luteinising hormone release during fasting in female rats. Journal of Neuroendocrinology 11, 429-434. Butler, J., Sjöberg, M. & Coen, C.W. (1999). Evidence for oestrogen receptor alphaimmunoreactivity in gonadotrophin-releasing hormone expressing neurons. Journal of Neuroendocrinology 11, 331-335. Butler, J., Kalló, I., Sjöberg, M. & Coen, C.W. (1999). Evidence for extensive distribution of estrogen receptor alpha-immunoreactivity in the cerebral cortex of adult rats. Journal of Neuroendocrinology 11 325-329. Cagampang, F.R.A., Piggins, H.D., Sheward, W.J., Harmar, A.J. & Coen, C.W. (1998). Circadian changes in PACAP type 1 (PAC1) receptor mRNA in the rat suprachiasmatic and supraoptic nuclei. Brain Research 813, 218-222. Cagampang, F.R.A., Antoni, F.A., Smith, S.M., Piggins, H.D. & Coen, C.W. (1998). Circadian changes of type II adenylyl cyclase mRNA in the rat suprachiasmatic nuclei. Brain Research 810, 279-282. Lucas, R.J., Cagampang F.R.A., Loudon, A.S.I., Stirland, J.A. & Coen, C.W. (1998). Expression of vasoactive intestinal peptide mRNA in the suprachiasmatic nuclei of the circadian tau mutant hamster. Neuroscience Letters 249, 147-150. Kalló, I., Fekete, C., Coen, C.W. & Liposits Z. (1998). Synaptic connections between substance P-containing axons and estrogen receptor-synthesizing neurons in the medial preoptic area of the rat brain. Brain Research 781, 335-338. Cagampang, F.R.A., Sheward, W.J., Harmar, A.J., Piggins, H.D. & Coen, C.W. (1998). Circadian changes in the expression of vasoactive intestinal peptide 2 receptor mRNA in the rat suprachiasmatic nuclei. Molecular Brain Research 54, 108-112. Cagampang, F.R.A., Rattray, M., Powell, J.F., Campbell, I.C. & Coen, C.W. (1998). Variation in the expression of the mRNA for protein kinase C isoforms in the rat suprachiasmatic nuclei, caudate putamen and cerebral cortex. Molecular Brain Research 53, 277-284. Cagampang, F.R.A., Cates, P.S., Sandhu, S., Strutton, P.H., McGarvey, C., Coen, C.W. & O’Byrne, K.T. (1997). Hypoglycaemia-induced inhibition of pulsatile luteinising hormone 5 secretion in female rats: role of oestradiol, endogenous opioids and the adrenal medulla. Journal of Neuroendocrinology 9, 867-872. Coen, C.W., Strutton, P.H., Cagampang, F.R.A., Butler, J.A. & O’Byrne, K.T. (1997). Investigating the luteinizing hormone pulse generator in rats: the importance of lateral thinking. In: Neural Control of Reproduction: Physiology and Behavior, pp. 85-102. Eds. A. Yokoyama, K-I. Maeda & H. Tsukamura: Japan Scientific Societies Press, Tokyo. Strutton, P.H. & Coen, C.W. (1996). Sodium pentobarbitone and the suppression of luteinizing hormone pulses in the female rat: the role of hypothermia. Journal of Neuroendocrinology 8, 941-946. Kalló I., Fekete C., Coen C.W., Flerkó B. & Liposits Z. (1996). Immunocytochemical characterization of the estrogen-receptive neurons and their afferent connections in the rodent brain. Neurobiology 4, 330-336. Cagampang, F.R.A., Rattray, M., Powell, J.F., Campbell, I.C. & Coen, C.W. (1996). Circadian changes of glutamate decarboxylase 65 and 67 mRNA in the rat suprachiasmatic nuclei. NeuroReport 7, 1925-1928. Chong, N.W.S., Cagampang, F.R.A., Coen, C.W., Campbell, I.C. & Powell, J.F. (1996). Rapid identification of novel genes expressed in a circadian manner in rat suprachiasmatic nuclei. NeuroReport 7, 1199-1203. Cagampang, F.R.A., Rattray, M., Powell, J.F., Chong, N.W.S., Campbell, I.C. & Coen, C.W. (1996). Circadian variation of EAAC1 glutamate transporter messenger RNA in the rat suprachiasmatic nuclei. Molecular Brain Research 35, 190-196. Persaud, R.A.P. & Coen, C.W. (1994). Effects of bombesin on core temperature in rats: studies involving radiotelemetry. NeuroReport 5, 841-843. Maeda, K-I., Cagampang, F.R.A., Coen, C.W. & Tsukamura, H. (1994). Involvement of the catecholaminergic input to the paraventricular nucleus and of corticotrophin-releasing hormone in the fasting-induced suppression of luteinizing hormone release in female rats. Endocrinology 134, 1718-1722. Clarke, I.J., Jessop, D.S., Millar, R.P., Morris, M., Bloom, S.R., Lightman, S.L., Coen, C.W., Lew, R.A. & Smith, A.I. (1993). Many peptides that are present in the external zone of the median eminence are not secreted into the hypophysial portal blood of sheep. Neuroendocrinology 57, 765-775. Tsukamura, H., Ohkura, S., Coen, C.W. & Maeda, K-I. (1993). The paraventricular nucleus and corticotrophin-releasing hormone are not critical in suppressing pulsatile LH secretion in ovariectomized lactating rats. Journal of Endocrinology 137, 291-297. Coen, C.W. (1993). The central control of luteinizing hormone release: a provisional model. In: Endocrine and Nutritional Control of Basic Biological Functions, pp. 441-454. Eds. H. Lehnert, R. Murison, H. Weiner, D. Hellhammer and J. Beyer: Hogrefe & Huber Publishers. Kalló, I., Coen, C.W., Fekete, C., Flerkó, B. & Liposits, Z. (1992). Relationship between galanin and estrogen receptor containing neuronal structures as revealed by immunocytochemistry. The Histochemical Journal 24, 18-19. Cagampang, F.R.A., Ohkura, S., Tsukamura, H., Coen, C.W., Ota, K. & Maeda, K-I. (1992). α2Adrenergic receptors are involved in the suppression of luteinizing hormone 6 release during acute fasting in the ovariectomized estradiol-primed rats. Neuroendocrinology 56, 724-728. Kalló, I., Liposits, Z., Flerkó, B. & Coen, C.W. (1992). Immunocytochemical characterization of afferents to estrogen receptor containing neurons in the medial preoptic area of the rat. Neuroscience 50, 299-308. Flerkó, B., Kalló, I., Coen, C.W. & Liposits, Z. (1992). Estrogen-sensitive neurons in the forebrain of the rat: from hypothesis to ultrastructural localization. New Developments in Biosciences 6, 37-56. Castaneyra-Perdomo, A., Perez-Delgado, M.M., Montagnese, C. & Coen, C.W. (1992). Brainstem projections to the medial preoptic region containing the luteinizing hormone-releasing hormone perikarya in the rat: an immunohistochemical and retrograde transport study. Neuroscience Letters 139, 135-139. Ruiz de Elvira, M.C., Persaud, R. & Coen, C.W. (1992). Use of running-wheels regulates the effects of the ovaries on circadian rhythms in rats. Physiology and Behavior 52, 277-284. Flerkó, B., Kalló, I., Coen, C.W. & Liposits, Z. (1991). Estrogen sensitive neurons in the hypothalamus of the rat. In: The Neuroendocrinology of Sex Steroids: Basic Knowledge, Clinical Applications, pp. 101-114. Opacka-Juffry, J., Ruiz de Elvira, M.C., Tacconelli, F. & Coen, C.W. (1991). Catecholamine synthesis inhibitors increase pineal adrenaline content by stimulating adrenal medullary activity. Neuroscience 42, 291-297. Ruiz de Elvira, M.C. & Coen, C.W. (1990). Centrally administered Neuropeptide Y enhances the hypothermia induced by peripheral administration of adrenoceptor antagonists. Peptides 11, 963967. O’Byrne, K.T., Lunn, S.F. & Coen, C.W. (1990). Central oxytocin stimulates luteinizing hormone release in the marmoset, a primate which fails to show lactationally-induced infertility. Journal of Neuroendocrinology 2, 419-421. Coen, C.W., Montagnese, C. & Opacka-Juffry, J. (1990). Coexistence of gonadotrophin releasing hormone and galanin: immunohistochemical and functional studies. Journal of Neuroendocrinology 2, 107-111. Liposits, Z., Kalló, I., Coen, C.W., Paull, W.K. & Flerkó, B. (1990). Ultrastructural analysis of estrogen receptor immunoreactive neurons in the medial preoptic area of the female rat brain. Histochemistry 93, 233-239. Coen, C.W. (1988). Pharmacological and neuroanatomical studies on hypothalamic adrenaline and LH release. In: The Brain and Female Reproductive Function - Basic and Clinical Aspects, pp. 35-43. Eds. F. Petraglia and C. Nappi: Parthenon Publishing. Opacka-Juffry, J., Tacconelli, F. & Coen, C.W. (1988). Sensitive method for determination of picogram amounts of epinephrine and other catecholamines in microdissected samples of rat brain using liquid chromatography with electrochemical detection. Journal of Chromatography 433, 41-51. Coen, C.W. (1987). Hypothalamic biogenic amines and the regulation of luteinizing hormone release in the rat. In: Endocrinology and Physiology of Reproduction, pp. 71-98. Eds. P.C.K. Leung, D.T. Armstrong, K.B. Ruf, W. Moger and H.G. Friesen: Plenum Press.7 Coen, C.W. & Gallo, R.V. (1986). Effects of various inhibitors of phenylethanolamine Nmethyltransferase on pulsatile release of luteinizing hormone in ovariectomized rats. Journal of Endocrinology 111, 51-59. Coen, C.W., Simonyi, A. & Fekete, M.I.K. (1985). Effect of phenylethanolamine Nmethyltransferase inhibition on serum concentrations of luteinizing hormone, follicle stimulating hormone and prolactin in pro-oestrous and ovariectomized rats. Brain Research 343, 383-387. Coombs, M.C. & Coen, C.W. (1983). Adrenaline turnover rates in the medial preoptic area and mediobasal hypothalamus in relation to the release of luteinizing hormone in female rats. Neuroscience 10, 207-210. Coen, C.W. & Coombs, M.C. (1983). Effects of manipulating catecholamines on the incidence of the preovulatory surge of luteinizing hormone and ovulation in the rat: evidence for a necessary involvement of hypothalamic adrenaline in the normal or ‘midnight’ surge. Neuroscience 10, 187-206. Coen, C.W., Coombs, M.C., Wilson, P.M.J., Clement, E.M. & MacKinnon, P.C.B. (1983). Possible resolution of a paradox concerning the use of p-chlorophenylalanine and 5hydroxytryptophan: evidence for a mode of action involving adrenaline in manipulating the surge of luteinizing hormone in rats. Neuroscience 8, 583-591. Coen, C.W. & MacKinnon, P.C.B. (1981). Induction of ovulation in PCPA-blocked intact adult rats by 5-HTP: effect of PCPA on ovarian function. Advances in Physiological Sciences 14 (II), 185-190. Coen, C.W., Franklin, M., Laynes, R.W. & MacKinnon, P.C.B. (1980). Effects of manipulating serotonin on the incidence of ovulation in the rat. Journal of Endocrinology 87, 195-201. Coen, C.W. & MacKinnon, P.C.B. (1980). Lesions of the suprachiasmatic nuclei and the serotonin-dependent phasic release of luteinizing hormone in the rat: effects on drinking rhythmicity and on the consequences of preoptic area stimulation. Journal of Endocrinology 84, 231-236. Coen, C.W. & MacKinnon, P.C.B. (1979). Serotonin involvement in the control of phasic luteinizing hormone release in the rat: evidence for a critical period. Journal of Endocrinology 82, 105-113. Garrud, P., Gray, J.A., Rickwood, L. & Coen, C.W. (1977). Pituitary-adrenal hormones and effects of partial reinforcement on appetitive behaviour in the rat. Physiology and Behaviour 18, 813-818.