Flexibility in an emergency life-history stage: acute food deprivation prevents sickness behaviour but not the immune response.

The emergency life-history stage (ELHS) can be divided into two subcategories that describe distinct, coordinated responses to disease- or non-disease-related physiological challenges. Whether an individual can simultaneously express aspects of both subcategories when faced with multiple challenges is poorly understood. Emergency life-history theory suggests that disease- and non-disease-related responses are coordinated at the level of the whole organism and therefore cannot be expressed simultaneously. However, the reactive scope and physiological regulatory network models suggest that traits can be independently regulated, allowing for components of both disease- and non-disease-related responses to be simultaneously expressed within a single organism. To test these ideas experimentally, we subjected female zebra finches to food deprivation, an immune challenge, both, or neither, and measured a suite of behavioural and physiological traits involved in the ELHS. We examined whether the trait values expressed by birds experiencing simultaneous challenges resembled trait values of birds experiencing a single challenge or if birds could express a mixture of trait values concurrently. We find that birds can respond to simultaneous challenges by regulating components of the behavioural and immune responses independently of one another. Modularity within these physio-behavioural networks adds additional dimensions to how we evaluate the intensity or quality of an ELHS. Whether modularity provides fitness advantages or costs in nature remains to be determined.

Neural Versus Gonadal GnIH: Are they Independent Systems? A Mini-Review.

Based on research in protochordates and basal vertebrates, we know that communication across the first endocrine axes likely relied on diffusion. Because diffusion is relatively slow, rapid responses to some cues, including stress-related cues, may have required further local control of axis outputs (e.g., steroid hormone production by the gonads). Despite the evolution of much more efficient circulatory systems and complex nervous systems in vertebrates, production of many "neuro"transmitters has been identified outside of the hypothalamus across the vertebrate phylogeny and these neurotransmitters are known to locally regulate endocrine function. Our understanding of tissue-specific neuropeptide expression and their role coordinating physiological/behavioral responses of the whole organism remains limited, in part, due to nomenclature and historic dogma that ignores local regulation of axis output. Here, we review regulation of gonadotropin-inhibitory hormone (GnIH) across the reproductive axis in birds and mammals to bring further attention to context-dependent disparities and similarities in neuropeptide production by the brain and gonads. We find that GnIH responsiveness to cues of stress appears conserved across species, but that the response of specific tissues and the direction of GnIH regulation varies. The implications of differential regulation across tissues remain unclear in most studies, but further work that manipulates and contrasts function in different tissues has the potential to inform us about both organism-specific function and endocrine axis evolution.

Neural and neuroendocrine processing of a non-photic cue in an opportunistically breeding songbird.

Recent studies of the onset of breeding in long-day photoperiodic breeders have focused on the roles of type 2 and 3 iodothyronine deiodinases (DIO2 and DIO3) in the conversion of thyroxine (T4) to triiodothyronine (T3) and subsequent activation of the reproductive axis. It has been hypothesized that an increase in DIO2 and a reciprocal decrease in DIO3 causes the release of gonadotropin-releasing hormone (GnRH) from the hypothalamus, setting off a reproductive cascade, and that this DIO mechanism for GnRH release is conserved across vertebrate taxa. We sought to test whether social cues that are known to stimulate reproductive behaviors can activate the DIO system to initiate reproduction in a non-photoperiodic bird, the zebra finch (Taeniopygia guttata). Isolation of males and subsequent presentation of females did not increase DIO2 or GnRH expression in the hypothalamus, nor did it decrease gonadotropin-inhibitory hormone (GnIH) or DIO3. Males receiving a female stimulus showed significantly higher mRNA expression and immunoreactive cell count of the immediate-early gene early growth response protein 1 (EGR-1) than isolated males, indicating hypothalamic activation in response to a female. Cells immunoreactive for EGR-1 were not co-localized with those immunoreactive for GnRH. Reproductive behaviors (singing, copulation attempts and overall activity) were significantly higher in males receiving a female stimulus. This study presents a social effect on behavior and EGR-1 expression in the hypothalamus of males in response to females, but more research is needed to determine whether the DIO2 system and the GnRH system are responsive to social stimulation in this species.

Differential response of GnIH in the brain and gonads following acute stress in a songbird.

Gonadotropin-inhibitory hormone (GnIH) acts to inhibit reproduction at all levels of the hypothalamo-pituitary-gonad axis. GnIH expression and/or immunoreactivity in the hypothalamus increase with acute stress in some birds and mammals, and thus may be involved in stress-induced reproductive inhibition. Much is known about GnIH and stress in seasonal and continuous breeders, but far less is known about these interactions in opportunistic breeders. For opportunistically breeding animals, reproductive readiness is closely associated with unpredictable environmental cues, and thus the GnIH system may be more sensitive to stress. To test this, we collected tissues from zebra finches immediately following capture or after 60 min of restraint. Restraint significantly increased plasma corticosterone in males and females but, contrary to studies on other species, restrained birds had significantly fewer GnIH immunoreactive (GnIH-ir) cell bodies than control birds. GnIH-ir cell number did not differ between the sexes. Stressed females had lower mRNA expression of the beta subunit of follicle stimulating hormone (FSHß) in the pituitary, suggesting that the reduction in observed GnIH immunoreactivity in females may have been due to increased GnIH release in response to acute stress. GnIH expression increased in the testes, but not the ovaries, of restrained animals. Our data suggest that although GnIH responsiveness to stress appears to be conserved across species, specific tissue response and direction of GnIH regulation is not. Variation in the GnIH response to stress between species might be the result of ecological adaptations or other species differences in the response of the GnIH system to stress.