Abstract

Urbanization is a critical form of environmental change that can affect the physiology and behavior of wild animals and, notably, birds. One behavioral difference between birds living in urban and rural habitats is that urban males show elevated boldness or territorial aggression in response to simulated social challenge. This pattern has been described in several populations of song sparrow, Melospiza melodia. Such behavioral differences must be underpinned by differences in the brain, yet little work has explored how urbanization and neural function may be interrelated. We explored the relationship between urbanization and neural activation within a network of brain regions, collectively called the social behavior network, which contributes to the regulation of territorial aggression. Specifically, we captured free-living, territorial male song sparrows by playing them conspecific songs for 6-11 minutes, and then collected their brains. We estimated recent neural activation, as indicated by the immediate early gene FOS, and measured levels of a neuropeptide, arginine vasotocin (AVT), which is involved in the regulation of social behavior. Based on previous studies we expected urban males, which are generally more territorially aggressive, to have lower FOS expression in a node of the social behavior network implicated in regulating territoriality, the lateral septum (LS). Additionally, we expected urban males to have lower AVT expression in a brain region involved in the regulation of sociality, the medial bed nucleus of the stria terminalis (BSTm). We found that, compared to rural males, urban male song sparrows did have lower FOS expression in the LS. This pattern suggests that lower neural activation in the LS could contribute to behavioral adjustments to urbanization in male song sparrows. Additionally, counter to our predictions, urban male song sparrows had higher AVT-like immunoreactivity in the BSTm. Future work building upon these findings is needed to determine the causal role of such neural differences across rural and urban habitats. Understanding the mechanisms impacted by urbanization will inform our understanding of the reversibility and consequences of this form of habitat change.

Highlights

  • There is an urgent need to understand the endocrine and neuroendocrine mechanisms that permit animals to cope with changing environments because elucidating how animals adjust their physiology to urbanization will shed light on why some species persist and others decline when faced with a changing environment (Cockrem, 2005; Visser, 2008; Wingfield, 2008; Whitman and Agrawal, 2009; Engel et al, 2011; Hoffmann and Sgrò, 2011; Wong and Candolin, 2015)

  • Though there was a trend for the day of sampling to impact plasma testosterone such that testosterone was counterintuitively higher later in the season (GLM, effect of day of year, t = 2.093, P = 0.058), there were no overall differences in testosterone levels as a function of habitat type when day of year was taken into account

  • We found a significant effect of habitat type on FOS-ir in the LSc.vl (GLMM, habitat x LSc.vl, t = −3.105, P = 0.013) and, nearly, in the bed nucleus of the stria terminalis (BSTm) (GLMM, habitat x BSTm, t = −2.174, P = 0.058; Table 1; Figure 3)

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Summary

Introduction

Anthropogenic habitat disturbance is recognized as impacting the phenotypes of wild animals and is a particular concern for wild birds (Crick, 2004; Both et al, 2006; Caro, 2007; Visser, 2008; Wingfield, 2008; Bonier, 2012; Sol et al, 2013; Wong and Candolin, 2015). Urbanization Impacts Song Sparrow’s Brains and physiology through phenotypic plasticity to cope with such environmental change (Vitousek et al, 1997; Wingfield, 2008; Bonier, 2012; Sol et al, 2013; Wong and Candolin, 2015). Increased boldness or aggression in urban male song sparrows could be a response to resource availability (Foltz et al, 2015b) or differences in conspecific density (though see Davies and Sewall, 2016). Despite repeated demonstrations of behavioral differences in song sparrows living along urban-rural gradients, the neural and physiological basis of differences in aggression are not fully understood (Evans et al, 2010; Foltz et al, 2015a; Davies and Sewall, 2016)

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