Short day lengths alter stress and depressive-like responses, and hippocampal
morphology in Siberian hamsters
Joanna L. Workman
a,
⁎, Natalie Manny
a
, James C. Walton
b
, Randy J. Nelson
a, b, c
a
Department of Psychology, The Ohio State University, Columbus, OH 43201, USA
b
Department of Neuroscience, The Ohio State University, Columbus, OH 43201, USA
c
Institute for Behavioral Medicine Research, The Ohio State University, Columbus, OH 43201, USA
abstract article info
Article history:
Received 5 April 2011
Revised 29 July 2011
Accepted 31 July 2011
Available online 7 August 2011
Keywords:
Hippocampus
Depression
Stress
Photoperiod
Seasonality
Plasticity
Seasonal affective disorder
Many psychological disorders comprise a seasonal component. For instance, seasonal affective disorder
(SAD) is characterized by depression during autumn and winter. Because hippocampal atrophy may
underlie the symptoms of depression and depressive-like behaviors, one goal of this study was to determine
whether short days also induce structural changes in the hippocampus using photoperiod responsive
rodents — Siberian hamsters. Exposure to short days increases depressive-like responses (increased
immobility in the forced swim test) in hamsters. Male hamsters were housed in either short (LD 8:16) or
long days (LD 16:8) for 10 weeks and tested in the forced swim test. Brains were removed and processed for
Golgi impregnation. HPA axis function may account for photoperiod-related changes in depressive-like
responses. Thus, stress reactivity was assessed in another cohort of photoperiod-manipulated animals. Short
days reduced soma size and dendritic complexity in the CA1 region. Photoperiod did not induce gross
changes in stress reactivity, but an acute stressor disrupted the typical nocturnal peak in cortisol
concentrations. These data reveal that immobility induced by exposure to short days is correlated with
reduced CA1 cell complexity (and perhaps connectivity). This study is the first to investigate hippocampal
changes in the context of short-day induced immobility and may be relevant for understanding
psychological disorders with a seasonal component.
© 2011 Elsevier Inc. All rights reserved.
Introduction
In seasonally-changing environments, small mammals must alter
physiology and behavior in order to coordinate endogenous processes
with ambient conditions. Winter is a particularly challenging time to
survive and reproduce. In order to reliably coordinate physiological
processes with environmental conditions, many rodents monitor day
length, a precise and relatively noise-free cue. Day length (photope-
riod) information is encoded physiologically through the duration
of nighttime melatonin secretion, which is inversely proportional to
day length (Reiter, 1993). In addition to coordinating reproductive
processes, day length alters many nonreproductive behavioral and
physiological changes. Seasonally-breeding rodents, for instance,
undergo changes in brain morphology dependent on photoperiod.
Wild-caught rodents have reduced skull size (Pucek, 1963), whole
brain weights and hippocampal weight in winter (Yaskin, 1984).
Exposure to short days in the laboratory reduces whole brain and
hippocampal volume in white-footed mice (Peromyscus leucopus)
(Perrot-Sinal et al., 1998; Pyter et al., 2005) suggesting that changes in
brain volume in the wild may reflect changes induced by decreased
day length. The brain requires a disproportionate amount of energy
compared to other tissues, consequently slight reductions in brain
volume may be adaptive because they help conserve energy in harsh
environments. Seasonally-breeding rodents can be used in the
laboratory to isolate the effects of day length on brain morphology;
research in this area may reveal mechanisms of how day length alters
the brain (and subsequently mental processes) in humans.
SAD is one disorder with a strong seasonal component, and is a
subtype of major depressive disorder that encompasses similar
symptoms to major depression (e.g., depressed mood, anhedonia,
guilt), but some atypical symptoms as well (e.g., hypersomnia,
hyperphagia). These symptoms remit in the summer and some
individuals may become hypomanic. SAD has been associated with
an inability to inhibit morning melatonin secretion in the winter (Lewy
et al., 1999; Lewy and Sack, 1988; Rosenthal et al., 1984; Wehr et al.,
2001) suggesting that shortened day length, and in turn extended
melatonin secretion, may induce symptoms in vulnerable individuals.
Currently, there is no animal model of SAD but several species exhibit
greater depressive-like responses after exposure to short days. The
forced swim test (FST) is a well-validated behavioral measure used to
Hormones and Behavior 60 (2011) 520–528
⁎ Corresponding author at: The University of British Columbia, 2136 West Mall,
Vancouver, Canada BC V6T 1Z4.
E-mail address: joanna@psych.ubc.ca (J.L. Workman).
0018-506X/$ – see front matter © 2011 Elsevier Inc. All rights reserved.
doi:10.1016/j.yhbeh.2011.07.021
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