Sex differences, but no seasonal variations in the hippocampus of food-caching squirrels: a stereological study.

Recent studies have described sex differences in the relative size of the hippocampus that are associated with sex differences in space use in birds and short-lived mammals. A correlation between spatial learning and increased hippocampal volume has also been demonstrated in food-caching animals. Such results suggest that sexually dimorphic spatial learning (sex differences in space use during the breeding season) and seasonal variations in food-caching behavior (spatial memory for cache locations) might correlate with morphological changes in the hippocampus of adult long-lived mammals. We used modern stereological techniques to examine the volume and neuron number of the structures forming the hippocampal complex (dentate gyrus, CA3, and CA1) of wild adult eastern gray squirrels (Sciurus carolinensis) throughout the year. We observed differences in brain size between samples collected at different times of the year (October, January, and June). Our analysis showed sex differences, but no seasonal variations, in the volume of CA1 stratum oriens and stratum radiatum. There were no sex differences or seasonal variations in the relative volume or the number of neurons of any other layer of the structures forming the hippocampal complex. These results confirm the existence of sex differences in the structure of the hippocampus; however, this sexual dimorphism does not vary seasonally in adulthood and is likely to result from developmental processes. These results do not support the hypothesis that seasonal variations in food-caching behavior might correlate with morphological changes, such as variations in volume or neuron number, in the hippocampal complex of adult long-lived mammals.

The seasonal pattern of cell proliferation and neuron number in the dentate gyrus of wild adult eastern grey squirrels.

The dentate gyrus is one of two areas in the mammalian brain that produces neurons in adulthood. Neurogenesis (proliferation, survival, and differentiation of new neurons) is regulated by experience, and increased neurogenesis appears to be correlated with improved spatial learning in mammals and birds. We tested the hypothesis that in long-lived mammals that scatter-hoard food, seasonal variations in spatial memory processing (i.e. increased processing during caching season in the autumn) might correlate with changes in neurogenesis and neuron number in the granule cell layer of the dentate gyrus (gcl DG). We investigated the rate of cell proliferation and the total number of neurons in the granule cell layer of wild adult eastern grey squirrels (Sciurus carolinensis) at three different times of the year (October, January and June). We found no seasonal differences in cell proliferation rate or in total neuron number in the granule cell layer. Our findings are in agreement with those of previous studies in laboratory mice and rats, and in free-ranging, food-caching, black-capped chickadees, as well as with current hypotheses regarding the relationship between neurogenesis and learning. Our results, however, are also in agreement with the hypothesis that neurogenesis in the dentate gyrus represents a maintenance system that may be regulated by environmental factors, and that changes in total neuron number previously reported in rodents represent developmental changes rather than adult plasticity. The patterns observed in mature wild rodents, such as free-ranging squirrels, may represent more accurately the extent of hippocampal plasticity in adult mammals.

The economy of winter: phenotypic plasticity in behavior and brain structure.

Mobile animals must learn the spatial distributions of resources. The cost of foraging increases dramatically for temperate-zone animals during the winter. Two strategies may be used to balance the energetic budget: reducing costs of foraging and reducing need to forage. Both strategies are correlated with changes in brain structure, specifically in the hippocampus, a fore-brain structure used by birds and mammals to map spatial distributions of resources. Small mammals that reduce their need to forage, through hibernation or reduction in body size, show a specific reduction in the structure and size of the hippocampus. The costs of foraging can be also decreased by migration to better foraging conditions or by food-storing, both of which decrease the temporal heterogeneity of food resources. Both of these latter strategies are associated with increased hippocampal structure; for food-storing birds, this increase is a seasonal phenomenon. Thus not only behavior, but also learning ability and even brain structures in adult animals, may be phenotypically plastic in response to the changing demands of the environment.

Sexual selection and the brain.

Sex differences are intrinsically interesting, particularly in the brain. When sexually dimorphic structures mediate learning, and when such learning ability is necessary to compete for mates, then such differences are best understood within the framework of sexual selection. By categorizing recent studies of sex differences in the brain by their role in mate competition, theories of sexual selection can be used to predict and characterize the occurrence of dimorphisms among species with different mating systems.

Patient-controlled analgesia: a comparison of dosing regimens for acute postsurgical pain.

This study compares several dosing regimens for patient-controlled analgesia (PCA) in the management of acute maxillofacial surgical pain. The dosing methods differed by presence or absence of an active drug (morphine [MS] vs saline), presence of a baseline infusion, and dose of drug delivered. Sixty-eight patients were enrolled in this prospective, randomized, double-blind, placebo-controlled trial that lasted 24 hours. The study was completed in two separate parts, each of which involved randomization of patients into four groups (part I) or three groups (part II). No significant differences were noted in pain scores in the preoperative, immediate postoperative, or 4-, 8-, 12-, or 24-hour periods among any of the groups, including the saline-only control groups; in either part I or part II of the study. Significant differences (P < .01), however, were noted in nausea and vomiting scores. Fifty percent (50%) of patients receiving MS vomited, while no patients in the saline groups vomited. This study calls into question the usefulness of PCA with MS in maxillofacial surgery patients. Pain control was questionable at best, and the rate of emesis was unacceptably high in patients with potentially compromised airways. Further research is required to determine if other analgesics provide better pain control with less nausea in the PCA system or if antiemetics can effectively be used to lower the incidence of nausea and vomiting.

Natural space-use patterns and hippocampal size in kangaroo rats.

The size of the hippocampus, a forebrain structure that processes spatial information, correlates with the need to relocate food caches by passerine birds and with sex-specific patterns of space use in microtine rodents. The influences on hippocampal anatomy of sexual selection within species, and natural selection between species, have not yet been studied in concert, however. Here we report that natural space-use patterns predict hippocampal size within and between two species of kangaroo rats (Dipodomys). Differences in foraging behavior suggest that Merriam's kangaroo rats (D. merriami) require better spatial abilities than bannertail kangaroo rats (D. spectabilis). Sex-specific differences in mating strategy suggest that males of both species require more spatial ability than females. As predicted, hippocampal size (relative to brain size) is larger in Merriam's than in bannertail kangaroo rats, and males have larger hippocampi than females in both species. Males of a third species (D. ordii) also have smaller hippocampi than Merriam's kangaroo rat males, despite being similar to Merriam's in brain and body size. These results suggest that both natural and sexual selection affect the relative size and perhaps function of mammalian hippocampi. They also reassert that measures of functional subunits of the brain reveal more about brain evolution than measures of total brain size.

Spatial memory and adaptive specialization of the hippocampus.

The hippocampus plays an important role in spatial memory and spatial cognition in birds and mammals. Natural selection, sexual selection and artificial selection have resulted in an increase in the size of the hippocampus in a remarkably diverse group of animals that rely on spatial abilities to solve ecologically important problems. Food-storing birds remember the locations of large numbers of scattered caches. Polygynous male voles traverse large home ranges in search of mates. Kangaroo rats both cache food and exhibit a sex difference in home range size. In all of these species, an increase in the size of the hippocampus is associated with superior spatial ability. Artificial selection for homing ability has produced a comparable increase in the size of the hippocampus in homing pigeons, compared with other strains of domestic pigeon. Despite differences among these animals in their histories of selection and the genetic backgrounds on which selection has acted, there is a common relationship between relative hippocampal size and spatial ability.

Evolution of spatial cognition: sex-specific patterns of spatial behavior predict hippocampal size.

In a study of two congeneric rodent species, sex differences in hippocampal size were predicted by sex-specific patterns of spatial cognition. Hippocampal size is known to correlate positively with maze performance in laboratory mouse strains and with selective pressure for spatial memory among passerine bird species. In polygamous vole species (Rodentia: Microtus), males range more widely than females in the field and perform better on laboratory measures of spatial ability; both of these differences are absent in monogamous vole species. Ten females and males were taken from natural populations of two vole species, the polygamous meadow vole, M. pennsylvanicus, and the monogamous pine vole, M. pinetorum. Only in the polygamous species do males have larger hippocampi relative to the entire brain than do females. Two-way analysis of variance shows that the ratio of hippocampal volume to brain volume is differently related to sex in these two species. To our knowledge, no previous studies of hippocampal size have linked both evolutionary and psychometric data to hippocampal dimensions. Our controlled comparison suggests that evolution can produce adaptive sex differences in behavior and its neural substrate.

Regional differences in normally occurring cell death in the developing hamster lateral geniculate nuclei.

Normal cellular degeneration occurs in the lateral geniculate nuclei (LGN) of the hamster thalamus early in postnatal development. Degenerative debris can be observed in the ventral and dorsal nuclei at postnatal days 2-10 and is present in greater and more variable amounts in the ventral nucleus. Cell degeneration in the dorsal LGN is maximal at postnatal day 5, identical to the degeneration pattern of the hamster retina and superior colliculus, but shows a second peak at postnatal day 8 which may relate to the establishment of cortical connectivity. The incidence of degenerative debris is significantly higher in the peripheral margins of the dorsal nucleus, a pattern also seen in the retina and the superior colliculus, suggesting that a differential cell death may be involved in the formation of regional specializations in the visual system.