Both food-storing behaviour and the hippocampus change annually in food-storing birds. Food storing increases substantially in autumn and winter in chickadees and tits, jays and nutcrackers and nuthatches. The total size of the chickadee hippocampus increases in autumn and winter as does the rate of hippocampal neurogenesis. The hippocampus is necessary for accurate cache retrieval in food-storing birds and is much larger in food-storing birds than in non-storing passerines. It therefore seems probable that seasonal change in caching and seasonal change in the hippocampus are causally related. The peak in recruitment of new neurons into the hippocampus occurs before birds have completed food storing and cache retrieval for the year and may therefore be associated with spacing caches, encoding the spatial locations of caches, or creating a neuronal architecture involved in the recollection of cache sites. The factors controlling hippocampal plasticity in food-storing birds are not well understood. Photoperiodic manipulations that produce change in food-storing behaviour have no effect on either hippocampal size or neuronal recruitment. Available evidence suggests that changes in hippocampal size and neurogenesis may be a consequence of the behavioural and cognitive involvement of the hippocampus in storing and retrieving food.
Neuronal recruitment in the black-capped chickadee (Poecile atricapillus) hippocampus occurs at a higher rate in the fall than at other times of the year. As a means of determining whether this increase in recruitment results from greater neuron production, chickadees were caught in the wild between October and March and injected with the cell-birth marker 5-bromo-2Ј-deoxyuridine. Two weeks later, birds were killed by overdose, and hippocampal neuron production, apoptosis, neuron number, and hippocampal volume were determined. Chickadees collected in October, November, January, February, and March did not differ in neuron production, apoptosis, hippocampal volume, or neuron number. These findings indicate that increases in neuronal recruitment in the chickadee hippocampus in the fall do not result from increased neuron production, but instead, enhanced survival of new neurons.
Previous research has shown heightened recruitment of new neurons to the chickadee hippocampus in the fall. The present study was conducted to determine whether heightened fall recruitment is associated with the seasonal onset of food-storing by comparing neurogenesis in chickadees and a non-food-storing species, the house sparrow. Chickadees and house sparrows were captured in the wild in fall and spring and received multiple injections of the cell birth marker bromodeoxyuridine (BrdU). Birds were held in captivity and the level of hippocampal neuron recruitment was assessed after 6 weeks. Chickadees showed significantly more hippocampal neuronal recruitment than house sparrows. We found no seasonal differences in hippocampal neuronal recruitment in either species. In chickadees and in house sparrows, one-third of new cells labeled for BrdU also expressed the mature neuronal protein, NeuN. In a region adjacent to the hippocampus, the hyperpallium apicale, we observed no significant differences in neuronal recruitment between species or between seasons. Hippocampal volume and total neuron number both were greater in spring than in fall in chickadees, but no seasonal differences were observed in house sparrows. Enhanced neuronal recruitment in the hippocampus of food-storing chickadees suggests a degree of neurogenic specialization that may be associated with the spatial memory requirements of food-storing behavior.
Previous research presents a mixed picture of seasonal variation in the hippocampus of food-storing black-capped chickadees. One field study has shown an October peak in hippocampus volume, although laboratory studies conducted to determine whether photoperiod regulates this seasonal growth have failed to find changes in the size of the hippocampus. To resolve the discrepancy between field and lab reports we examined caching activity, hippocampal volume, and neurogenesis in adult male black-capped chickadees at four times over the annual cycle: October, January, April and July. We found that more birds stored food in October than at other times of year, but did not observe a significant change in the size of the hippocampus over the annual cycle. Telencephalon volume, however, was larger in October than in July. Hippocampal neuronal recruitment showed a significant peak in January, but there was no seasonal change in neuronal recruitment in the adjacent hyperpallium apicale. These results indicate that there might be seasonal variation in the recruitment of new neurons into the hippocampus of chickadees without overall seasonal change in hippocampal size.
Black-capped chickadees have a rich vocal repertoire including learned calls and the learned fee-bee song. However, the neural regions underlying these vocalizations, such as HVC, area X, and RA (robust nucleus of arcopallium), remain understudied. Here, we document seasonal changes in fee-bee song production and show a marked peak in singing rate during March through May. Despite this, we found only minimal seasonal plasticity in vocal control regions of the brain in males. There was no significant effect of time of year on the size of HVC, X, or RA in birds collected in January, April, July, and October. We then pooled birds into two groups, those with large testes (breeding condition) and those with small testes (nonbreeding), regardless of time of year. Breeding birds had slightly larger RA, but not HVC or X, than nonbreeding birds. Breeding birds had slightly larger HVC and RA, but not X, as a proportion of telencephalon volume than did nonbreeding birds. Birds collected in July had heavier brains than birds at other times of year, and had the greatest loss in brain mass during cryoprotection. The absence of any overall seasonal change in the vocal-control regions of chickadees likely results from a combination of individual differences in the timing of breeding phenology and demands on the vocal-control regions to produce learned calls year-round.
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