Abstract. Exposure of females to the male pheromone induces pulsatile release of gonadotropin-releasing hormone (GnRH) in goats. Recently, kisspeptin neurons in the arcuate nucleus (ARC) have been suggested to represent the proximate source of the GnRH pulse generator. In this study, we examined the effects of the pheromone on multipleunit activity (MUA) in female goats fitted with recording electrodes aimed at the ARC kisspeptin neurons. In all eight goats, periodic bursts in MUA (MUA volleys), which were considered to be electrophysiological manifestations of the GnRH pulse generator, were observed. The mean intervolley interval (T) during the control period was calculated in each goat that was then exposed to the male pheromone for 1 sec at timings of 1/4 T, 1/2 T or 3/4 T after one regularly occurring MUA volley. An instantaneous rise in MUA was observed immediately after the exposure regardless of timing. Exposure at a timing of 3/4 T resulted in an MUA volley within 60 sec following the instantaneous rise in all goats. In contrast, an MUA volley was induced in only 2 goats by exposure at 1/2 T, while exposure at 1/4 T failed to induce an MUA volley in any goats. These results suggest that transmission of the pheromone signal to the ARC, represented by an instantaneous rise, activates the GnRH pulse generator. Moreover, the timing-dependent pheromone action in inducing an MUA volley indicates that the GnRH pulse generator has a refractory period for the pheromone signal after the burst. Key words: GnRH pulse generator, Goat, Kisspeptin, Male effect, Pheromone (J. Reprod. Dev. 57 : 197-202, 2011) n goats and sheep, exposure of seasonally anestrous females to the primer pheromone secreted by sexually mature males results in an out-of-seasonal ovulation [1][2][3][4]. Since the initial endocrine event following reception of the pheromone is the stimulation of pulsatile luteinizing hormone (LH) secretion, it has been thought that the central target of the pheromone signal is the GnRH pulse generator [1,[3][4][5], i.e., the neural substrate that generates intermittent GnRH discharges into the portal vessels, thereby regulating pulsatile LH secretion into the peripheral circulation [6,7].Although the neural identity of the GnRH pulse generator per se was unclear, it was possible to measure its activity by recording multiple-unit activity (MUA) at the mediobasal hypothalamus. It has been demonstrated that electrophysiological manifestations of the GnRH pulse generator are represented as characteristic increases in MUA (MUA volleys) that are exclusively associated with LH pulses in the monkey [8], rat [9] and goat [10,11]. It might be that observed MUA volleys reflect the pulsatile activation of GnRH nerve terminals as they traverse en passant to the ME. Using this technique, Hamada et al. [12] showed that exposure of the female goat to male hair used as a pheromone source promptly induced an MUA volley, demonstrating for the first time that the pheromone action is in fact intimately associated with the GnRH pul...

Neurons in the arcuate nucleus (ARC) that concomitantly express kisspeptin, neurokinin B (NKB) and dynorphin A are termed KNDy neurons and are likely candidates for the intrinsic gonadotropin-releasing hormone (GnRH) pulse generator. Our hypothesis is that KNDy neurons are functionally and anatomically interconnected to generate discrete neural signals that govern pulsatile GnRH secretion. Our goal was to address this hypothesis using electrophysiological and anatomical experiments in goats. Bilateral electrodes targeting KNDy neurons were implanted into ovariectomized goats, and GnRH pulse generator activity, represented by characteristic increases in multiple-unit activity (MUA volleys), was measured. Spontaneous and pheromone- or senktide (an NKB receptor agonist)-induced MUA volleys were simultaneously recorded from both sides of the ARC. An anterograde tracer, biotinylated dextran amine (BDA), was also injected unilaterally into the ARC of castrated male goats, and the distribution of fibers containing both BDA and NKB was examined using dual-labeling histochemistry. The results showed that MUA volleys, regardless of origin (spontaneous or experimentally induced), occur simultaneously between the right and left sides of the ARC. Tract tracing indicated that axons projecting from NKB neurons in the ARC were directly apposed to other NKB neuronal cells located bilaterally in the ARC. These results demonstrate that GnRH pulse generator activity occurs synchronously between both sides of the ARC in goats and that KNDy neurons are bilaterally interconnected in the ARC via NKB-containing fibers. Taken together, the results suggest that KNDy neurons form a neuronal circuit to synchronize burst activity among KNDy neurons and thereby generate discrete neural signals that govern pulsatile GnRH secretion.

Collective behaviour is known to be the result of diverse dynamics and is sometimes likened to a living system. Although many studies have revealed the dynamics of various collective behaviours, their main focus was on the information process inside the collective, not on the whole system itself. For example, the qualitative difference between two elements and three elements as a system has rarely been investigated. Tononi et al. have proposed Integrated Information Theory (IIT) to measure the degree of consciousness Φ. IIT postulates that the amount of information loss caused by certain partitions is equivalent to the degree of information integration in the system. This measure is not only useful for estimating the degree of consciousness but can also be applied to more general network systems. Here we applied IIT (in particular, IIT 3.0 using PyPhi) to analyse real fish schools (Plecoglossus altivelis). Our hypothesis in this study is a very simple one: a living system evolves to raise its Φ value. If we accept this hypothesis, IIT reveals the existence of continuous and discontinuous properties as group size varies. For example, leadership in the fish school emerged for a school size of four or above; but not below three. Furthermore, this transition was not observed by measuring mutual information or in a simple Boids model. This result suggests that integrated information Φ can reveal some inherent properties which cannot be observed using other measures. We also discuss how the fish recognition of the figure-ground relation, that is, what determines the relevant ON and OFF states, may reveal various optimal paths for obtaining the functional evolution of collective behaviour.