Timothy Boswell

• PhD
• Newcastle University

Exposure to environmental stressors has detrimental phenotypic impacts on poultry welfare, but individual birds tend to respond differently. The density of highly plastic neurons in the hippocampus can serve as a biomarker of the experience of environmental stress in poultry, however, limited research has explored individual variation in this marke...

Recurring stress, which can be termed a negative cumulative affective experience has numerous detrimental phenotypic impacts on poultry welfare. Consequently, this negative experience could lead to reduced productivity. Biomarkers that serve as indicators of cumulative affective experience have been found through the density of highly plastic neuro...

It is still unclear which commercial housing system provides the best quality of life for laying hens. In addition, there are large individual differences in stress levels within a system. Hippocampal neurogenesis or plasticity may provide an integrated biomarker of the stressors experienced by an individual. We selected 12 adult hens each with goo...

It is still unclear which commercial housing system provides the best quality of life for laying hens. In addition, there are large individual differences in stress levels within a system. Hippocampal neurogenesis may provide an integrated biomarker of the stressors experienced by an individual. We selected 12 adult hens each with good and poor bod...

Broiler breeder chickens are commercially feed restricted to slow their growth and improve their health and production, however, there is research demonstrating that this leads to chronic hunger resulting in poor welfare. A challenge in these studies is to account for possible daily rhythms or the effects of time since last meal on measures relatin...

Access to outdoor areas is provided as a means of enhancing welfare in commercial systems for laying hens (Gallus gallus domesticus), but substantial individual differences exist in their proportional use. Baseline cell proliferation levels of Adult Hippocampal Neurogenesis (AHN) have been associated with individual differences in reactive vs. proa...

In commercial flocks of laying hens, keel bone fractures (KBFs) are prevalent and associated with behavioural indicators of pain. However, whether their impact is severe enough to induce a depressive-like state of chronic stress is unknown. As chronic stress downregulates adult hippocampal neurogenesis (AHN) in mammals and birds, we employ this mea...

In the mammalian brain, adult hippocampal neurogenesis (AHN) is suppressed by chronic stress, primarily at the ventral pole of the hippocampus. Based upon anatomy, we hypothesise that the caudal pole of the avian Hippocampal Formation (HF) presents a homologous subregion. We thus investigated whether AHN is preferentially suppressed in the caudal c...

Progress in improving the welfare of captive animals has been hindered by a lack of objective indicators to assess the quality of lifetime experience, often called cumulative affective experience. Recent developments in stress biology and psychiatry have shed new light on the role of the mammalian hippocampus in affective processes. Here we review...

Anselme & Güntürkün propose a novel mechanism to explain the increase in foraging motivation when experiencing an unpredictable food supply. However, the physiological mechanisms that maintain energy homeostasis already control foraging intensity in response to changes in energy balance. Therefore, unpredictability may just be one of many factors t...

Abstract Background Research into energy balance and growth has infrequently considered genetic sex, yet there is sexual dimorphism for growth across the animal kingdom. We test the hypothesis that in the chicken, there is a sex difference in arcuate nucleus neuropeptide gene expression, since previous research indicates hypothalamic AGRP expressio...

Background: Research into energy balance and growth has infrequently considered genetic sex, yet there is sexual dimorphism for growth across the animal kingdom. We test the hypothesis that sexual dimorphism for growth and body weight in birds is potentially dependent on hypothalamic agouti-related peptide (AGRP) expression, since previous research...

In birds little is knownabout the hormonal signals that communicate nutritional state to the brain and regulate appetitive behaviours. In mammals, the peptide hormones ghrelin and leptin elevate and inhibit consumption and food hoarding, respectively. But in birds, administration of both ghrelin and leptin inhibit food consumption. The role of thes...

The mammalian hippocampus is particularly vulnerable to chronic stress. Adult neurogenesis in the dentate gyrus is suppressed by chronic stress and by administration of glucocorticoid hormones. Post-natal and adult neurogenesis are present in the avian hippocampal formation as well, but much less is known about its sensitivity to chronic stressors....

Analysed data. This file contains all the data reported in the results section and presented in the figures. (XLSX)

Adult hippocampal neurogenesis (AHN) in the dentate gyrus is known to respond to environmental enrichment, chronic stress, and many other factors. The function of AHN may vary across the septo-temporal axis of the hippocampus, as different subdivisions are responsible for different functions. The dorsal pole regulates cognitive-related behaviors, w...

DG regions identification. To transfer the “X shaped structure” of pyramidal neuron belonging to CA3i (Fanselow and Dong, 2010) that denotes the iDG, we used images from the software Brain Explorer 2 (Allen Brain Institute, US). We initially define this region (A) and translate it from the sagittal to coronal plane (B). We then used these landmarks...

Thresholding method for DCX image quantification. Images acquired with 20X objective magnification were processed with ImageJ software and the GCL region (A) was outlined (black line) in all the analyzed frames. Images were acquired with the same amount of light and intra animal variability was overridden by setting the threshold level manually for...

PP-fold peptides such as peptide YY (PYY) and pancreatic polypeptide (PPY) are known to play key roles in vertebrate energy homeostasis. Until recently, no gene sequence was available for avian PYY and therefore a gap in knowledge of regulation of its expression exists in avian species. Here we further evidence the mRNA sequence for chicken PYY and...

The arcuate nucleus is generally conserved across vertebrate taxa in its neuroanatomy and neuropeptide expression. Gene expression of agouti-related protein (AGRP), neuropeptide Y (NPY), pro-opiomelanocortin (POMC), and cocaine- and amphetamine-regulated transcript (CART) has been established in the arcuate nucleus of several bird species and co-lo...

Animals can undergo periods of voluntarily reduced food intake and body weight or "animal anorexias" as part of their natural lives that are beneficial for survival or breeding such as during territorial behaviour, hibernation, migration and incubation of eggs. For incubation, a change in the defended level of body weight or "sliding set point" app...

The avian central melanocortin system is well conserved between birds and mammals in terms of the component genes, the localisation of their expression in the hypothalamic arcuate nucleus, the effects on feeding behaviour of their encoded peptides and the sensitivity of agouti-related protein (AGRP) and pro-opiomelanocortin (POMC) gene expression t...

The central melanocortin system is conserved across vertebrates. However, in birds, little is known about how energy balance influences orexigenic agouti-related protein (AGRP) and anorexigenic pro-opiomelanocortin (POMC) expression, despite the fact that commercial food restriction is critical to the efficient production of poultry meat. To enable...

Timothy Boswell trained in comparative endocrinology as an undergraduate and postgraduate student at the University of Bristol, England, in the laboratory of Brian Follett, graduating in 1991. He developed an interest in the seasonal regulation of food intake through his Ph.D. work on the regulation of migratory fattening in birds. Timothy pursued...

In order to further advance the understanding of genes involved in avian photoperiodic signaling, a chicken hypothalamic cDNA microarray was made to identify changes in gene expression in the whole hypothalamus of juvenile male domestic chickens after 4 days' photostimulation. The most robust change was a depression in heat shock protein 90B1 (HSP9...

Experimental studies of the physiological mechanisms underlying avian migration have concentrated on small passerines. The present study is concerned with the regulation of migratory fat deposition in a galliform. the European quail (Coturnix coturnix). The increased mass associated with migration was due exclusively to the deposition of fat wherea...

Endogenous plasma prolactin and baseline corticosterone concentrations were measured in Dark-eyed Juncos (Junco hyemalis, n=27) photostimulated into migratory condition to look at how these hormones may be linked to the development of migratory condition. In addition to the commonly used assay for corticosterone, a recombinant-derived European star...

Alpha-melanocyte-stimulating hormone (MSH) is well known as an anorexigenic peptide in the brain of mammals. In addition to this, brain alpha-MSH enhances heat production (HP), indicating that the peptide acts as a catabolic factor in the regulation of energy metabolism. The anorexigenic effect of alpha-MSH is also observed in chicks (Gallus gallus...

Glucagon-like peptide-1 (GLP-1), derived from proglucagon, is thought to act as a negative regulator of energy homeostasis in mammals, since intracerebroventricular (ICV) injection of GLP-1 inhibits feeding behavior and enhances energy expenditure. The anorexigenic effect of GLP-1 is also observed in chicks, but whether brain GLP-1 enhances energy...

Alpha-melanocyte-stimulating hormone (alpha-MSH) is recognized as an anorexic peptide in the brain of vertebrates, but its mechanism of action has not been identified in birds. Therefore, we investigated whether the anorexic effect of alpha-MSH is mediated by corticotrophin-releasing factor (CRF) in the domestic chick. Firstly, we found that intrac...

Vasoactive intestinal peptide (VIP) and pituitary adenylate cyclase-activating polypeptide (PACAP) are the members of the glucagon superfamily and bind to common receptors while PACAP also acts via the PACAP-specific receptor, PAC1. The aim of the present study was to investigate whether intracerebroventricular (i.c.v.) injection of VIP and PACAP a...

In mammals, alternative splicing of the leptin receptor (LEPR) produces several C-terminal truncated isoforms that are believed to play a role in the transport, cellular internalisation and degradation of the hormone leptin. The chicken leptin receptor (chLEPR) is similar to its mammalian counterparts in terms of its intron/exon structure and conse...

The purpose of the present study was to investigate whether intracerebroventricular (ICV) injection of neuropeptide Y (NPY) affects heat production (HP), body temperature, and plasma concentrations of metabolic fuels in chicks. ICV injection of NPY (0, 188 or 375 pmol) did not affect HP, but significantly lowered respiratory quotient as well as the...

The peptides of the neuropeptide Y (NPY) family exert their functions, including regulation of appetite and circadian rhythm, by binding to G-protein coupled receptors. Mammals have five subtypes, named Y1, Y2, Y4, Y5 and Y6, and recently Y7 has been discovered in fish and amphibians. In chicken we have previously characterized the first four subty...

Leptin is well established as a multifunctional cytokine in mammals. However, little is known about the evolution of the leptin gene in other vertebrates. A recently published set of ESTs from the tiger salamander (Ambystoma tigrinum) contains a sequence sharing 56% nucleotide sequence identity with the human leptin cDNA. To confirm that the EST is...

Prolactin-releasing peptide (PrRP) was named for its originally reported effects as a prolactin (PRL) secretagogue in mammals. Carassius RFamide (C-RFa) is an orthologous PRL secretagogue in fishes and a gene encoding a 20-amino acid peptide of identical sequence is present in the chicken. These facts suggest that C-RFa is a putative chicken PrRP....

The mammalian melanocortin system has been established as a crucial regulatory component in an extraordinarily diverse number of physiological functions. In contrast, comparatively little is known about the avian melanocortin system: interest in the physiological role of alpha-MSH in birds has been limited by the fact that birds lack the intermedia...

In domestic juvenile chickens kept on short days, photoinduced luteinsing hormone (LH) release, and by inference gonadotrophin-releasing hormone (GnRH) release, are readily detectable within 4 days of photostimulation.2 The molecular mechanisms responsible for the rapid photoinduced release of LH and GnRH in avian species are unknown. It has been s...

The physiological regulation of energy homeostasis is critical to an animal’s long-term survival and is relevant to the poultry industry in relation to animal production and welfare. In mammals, rapid progress has been made over the past decade in identifying the neural circuitry in the hypothalamus involved in the regulation of energy balance. Two...

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Studies performed in vitro suggest that a novel 12 amino acid RF amide peptide, isolated from the quail hypothalamus, is a gonadotrophin inhibitory hormone (GnIH). The aim of the present study was to investigate this hypothesis in the domestic chicken. Injections of GnIH into nest-deprived incubating hens failed to depress the concentration of plas...

Intracerebroventricular (ICV) injection of pituitary adenylate cyclase-activating polypeptide-38 (PACAP) or vasoactive intestinal peptide (VIP) inhibits feeding in chicks. However, the underlying anorexigenic mechanism(s) has not yet been investigated. The present study investigated whether these peptides influence the activity of corticotrophin-re...

Prolactin-releasing peptide (PrRP) is one of the inhibitory factors in feeding regulation of mammals. However, no information is available for avian species. The present study was done to clarify the effect of intracerebroventricular (ICV) injection of PrRP on feeding in chicks. Firstly, we found that ICV injection of PrRP (94-1500 pmol) significan...

Intracerebroventricular (ICV) injections of pituitary adenylate cyclase-activating polypeptide-38 (PACAP) and vasoactive intestinal peptide (VIP) inhibit feeding in chicks. However, the precise anorexigenic mechanisms have not been investigated, since both peptides activate the VPAC receptor in mammals. We investigated which receptor mediates the a...

Research in mammals has established the existence of a neuronal network that lies within the hypothalamus and that regulates energy homeostasis. However, it is unknown whether this system has been evolutionarily conserved. The objective of the present study was therefore to examine the influence of the agouti-related peptide (AGRP), pro-opiomelanoc...

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The interest in the physiological role of alpha-MSH in birds has been limited because they lack the intermediate lobe of the pituitary, the main source of circulating alpha-MSH in most vertebrates. Recent studies have improved our understanding of the avian melanocortin system. We have cloned and characterized all five MC-R subtypes, POMC, and AGRP...

The interest in the physiological role of α-MSH in birds has been limited because they lack the intermediate lobe of the pituitary, the main source of circulating α-MSH in most vertebrates. Recent studies have improved our understanding of the avian melanocortin system. We have cloned and characterized all five MC-R subtypes, POMC, and AGRP in chic...

Previous research has indicated an involvement of glucagon superfamily peptides in the regulation of feeding in the domestic chick brain. However the possible roles of vasoactive intestinal peptide (VIP) and pituitary adenylate cyclase-activating polypeptide-38 (PACAP) have not yet been investigated. We therefore examined the effect of intracerebro...

Chicken prepro-orexin cDNA has been cloned, sequenced and characterized. The predicted amino acid sequence of chicken prepro-orexin cDNA revealed that orexin-A and -B are highly conserved among vertebrate species. In situ hybridization and immunohistochemistry localized orexin-positive cell bodies in the periventricular hypothalamic nucleus extendi...

Within the neuropeptide Y (NPY) family of peptides, pancreatic polypeptide is the most divergent across species. It differs in 20 of 36 positions between human and chicken. In mammals, it binds primarily to the Y4 receptor, to which NPY and peptide YY (PYY) bind with lower affinities. Because of these large sequence differences in pancreatic polype...

The neuropeptide Y (NPY) receptor subtypes Y1 and Y5 are involved in the regulation of feeding and several other physiological functions in mammals. To increase our understanding of the origin and mechanisms of the complex NPY system, we report here the cloning and pharmacological characterization of receptors Y1 and Y5 in the first non-mammal, chi...

The neural mechanisms involved in the compensatory hyperphagia exhibited by many vertebrate species after a fast are not fully understood but, in mammals, appear to involve nutritionally-sensitive neurons that co-express neuropeptide Y (NPY) and agouti-related protein (AGRP) in the infundibular hypothalamus. We investigated whether these neurons ha...

The presence and possible physiological roles of alpha-melanocyte-stimulating hormone (alpha-MSH) in the peripheral tissues of birds have not been established. By a combination of RT-PCR, immunocytochemistry and in situ hybridization, we have examined alpha-MSH expression in the eye of the chicken during development. In the 1-day-old chick, alpha-M...

Here we report the molecular cloning of the chicken (Gallus gallus) neuropeptide Y (NPY) receptor Y2, the first non-mammalian Y2 receptor. It displays 75-80% identity to mammalian Y2 and has a surprisingly divergent cytoplasmic tail. Expression of the receptor protein in a cell line showed that the receptor did not bind the mammalian Y2 selective a...

Neuropeptides such as corticotropin-releasing factor (CRF) may play a role in regulating the pronounced seasonal changes in food intake shown by white-crowned sparrows (Zonotrichia leucophrys gambelii). White-crowned sparrows held on short day length received injections into the third ventricle (icv) of saline or 5.0, 15.0, and 30 microg/kg. Meal s...

The DNA sequence of a chicken leptin gene that shares 95% nucleotide similarity with the mouse leptin sequence has been recently reported (Taouis et al., 1998, Gene 208, 239-242). Experiments have been performed independently in two laboratories to try to confirm this finding. Fourteen PCR primers based on the mouse leptin sequence were designed to...

Lines of mice have been developed in our laboratory by divergent long-term selection for body fat content. This has resulted in a fivefold (23% vs 4%) higher fat percentage in the Fat line at 14 weeks of age, with little difference between the Fat and Lean lines in fat-free body weight. As part of an approach to characterize the physiological mecha...

As part of an approach to uncover the brain mechanisms underlying the regulation of energy balance in broiler chickens, we investigated the possible role of neuropeptide Y (NPY). The NPY gene expression was measured in the hypothalamus of birds from a standard Ross male line and a Ross relaxed line. Both lines are derived from the same founder stoc...

Systemic administration of cholecystokinin (CCK) inhibits feeding in birds. However, the signaling pathway through which CCK induces this effect is unknown, and its role as a natural satiety signal is controversial. To address these issues, we used immunocytochemistry for the immediate-early gene protein Fos to localize sites of neuronal activation...

Studies in vertebrates have shown that hormones can influence circadian rhythms of behavior. We investigated whether testosterone could induce rhythmicity in arrhythmic Japanese quail, kept in DD. The animals used were 31/2-week-old castrated males from a line of quail selected for the lack of the circadian rhythm of feeding activity. After 3 weeks...

Neuropeptide Y (NPY) has been implicated in the control of a number of physiological functions in birds including food intake and reproduction. In the present study, sites of NPY synthesis were localized in the brains of Japanese quail and domestic chickens by in situ hybridization histochemistry using a digoxigenin-labelled riboprobe. NPY mRNA was...

Migratory birds show a number of physiological adaptations for long distance flight, including increased storage and utilization of fat promoted by increased food intake before migration. However, the relationship between food intake and metabolic fuels in migratory birds has received little attention. To determine whether food intake is sensitive...

In changing environments, birds time reproduction to optimize survival of young (e.g., Wingfield, 1983; Perrins, 1970; Lack, 1968). Selective features of the environment (ultimate factors; Baker, 1938) favor those individuals that breed in reasonably close synchrony with the changes occurring in the environment. These ultimate factors include food...

Migratory birds rely on increased fat storage and fatty acid utilization to meet seasonal changes of energy expenditure and as a result increase food intake and fat stores before migration. To determine whether their feeding behavior is sensitive to carbohydrate and/or fatty acid utilization, white-crowned sparrows maintained on short daylength (9L...

Castration of certain passerine migrants on short days prevents fat deposition in the following spring. Previously, we showed that spring fattening is independent of the presence of the testes in European quail. However, a requirement for testosterone could not be ruled out. The aim of the present study was to compare the seasonal pattern of testos...

The present study addresses the role of prolactin as a regulator of migratory fattening in European quail ( Coturnix coturnix ). Plasma prolactin levels in captive birds undergoing migratory fattening in an outdoor aviary and in the laboratory were measured by radioimmunoassay with an antibody raised against recombinant-derived chicken prolactin. N...

To determine if altered sensitivity to neuropeptide Y (NPY) underlies premigratory fattening, white-crowned sparrows held on short day length (9:15-h light-dark) received injections into the third ventricle (ivt) of saline or several doses of NPY. An inverted-U function occurred with food intake increasing 30 and 60 min after doses of 1.0 and 2.0 m...

The aim of the present study was to determine the relationship between seasonal variation in levels of insulin, cortisol, and corticosterone and body mass in a natural population of golden-mantled ground squirrels (Spermophilus saturatus) over the active season, from April to August. The body mass of females is at an active season minimum on emerge...

White-crowned sparrows maintained on short days (9:15-h light-dark cycle) were peripherally injected with 1.0, 4.0, and 16 micrograms/kg ip of cholecystokinin octapeptide (CCK-8). Meal size over the subsequent 30 min was significantly depressed in a dose-dependent fashion. Water intake was not affected. The anorexic effect caused by 4.0 micrograms/...

This paper describes a method for chronically cannulating the third ventricle of the white-crowned sparrow, a small passerine bird, without damaging the midsagittal sinus. The method is reliable and chronic over at least 1 month. The technique was verified by assessing the effects of angiotensin II (ANG II) on inducing drinking behavior. All birds...

Neuropeptides such as neuropeptide Y (NPY) and galanin may play a role in regulating the pronounced seasonal changes in food intake shown by golden-mantled ground squirrels (Spermophilus saturatus). We used in situ hybridization histochemistry to localize the expression of NPY and galanin mRNA in the hypothalamus of normally feeding animals. NPY mR...