Literature Review of "Environmental experiences influence cortical volume in territorial

and nonterritorial side-blotched lizards, uta stansburiana" (LaDage et al., 2016)

Matt London
9/1/2017

Spatially based cognition, which is required for territoriality, navigation, and acquisition of
food resources, is regulated, in reptiles, by the medial cortex and dorsal cortex. The medial and
dorsal cortices are homologous to the hippocampus of mammals and are proposed to be
similarly involved in learning, memory, and navigation.Territorial males have larger cortices than
nonterritorial males, which is likely due to increased spatial processing demands for territorial
defense. It is unknown if cortical differences are result of genetics, spatial cognition, or
interaction of both. The authors proposed two hypotheses. One, that captivity deprives reptiles
of spatial processing demands and attenuates cortical volume and neurogenesis. And two,
correlational selection has linked morphotype/territorial predisposition (via the OBY locus) and
cortical volume, thus territorial and nonterritorial cortical differences will be stable across
environments. The authors predicted that laboratory-reared territorial males would have larger
cortices than laboratory reared non-territorial males and that laboratory-reared males would
exhibit smaller cortical volumes than wild-caught males.
Male side-blotched lizards (Uta stansburiana) were the reptiles utilized in this
experiment, comprising two phenotypes of territorial predisposition (orange and blue) and one
phenotype of nonterritorial disposition. The groups consisted of nine laboratory-reared territorial
(five orange and six blue) and nonterritorial (nine yellow) individually housed with 14:8 light:dark
cycle and ambient temperature between 25-40C. All lizards were sacrificed for brain analysis at
adulthood (10-12 months). Data from previous experiment were used to include wild-caught
territorial (nine orange and eight blue) and nonterritorial (seven yellow) collected from Los
Banos Grandes, CA. Since Uta stansburiana is annual species, they were between 9 and 12
months old when collected and sacrificed for brain analysis in March 2008. Brains were
sectioned in the coronal plane every 40 m, mounted, and Nissl stained with thionin. Medial and
Dorsal cortex sections were measured blind to territorial predisposition with standard
stereological methods. General linear models tested differences between territorial and
nonterritorial males in telencephalon to confirm that territoriality did not affect overall brain size.
A two-way MANOVA analyzed cortical volume with origin and territoriality as fixed factors and
telencephalon as a covariate (telencephalon minus cortical volumes).
Results revealed statistical significance in both cortical areas (medial and dorsal) for the
telencephalon covariate and origin (laboratory versus wild) of individuals. The interaction
between origin and territorial predisposition was only significant for the dorsal cortex. Compared
to wild-caught, territorial and non-territorial lab-reared males exhibited smaller medial and dorsal
cortices relative to remainder of telencephalon. Territorial wild-caught males had larger dorsal
cortices compared to the other three groups (nonterritorial wild-caught lizards and territorial and
nonterritorial laboratory reared males). This interaction between origin and territorial
predisposition was not significant between groups for the medial cortex. All lab-reared males
had similar cortical volume, regardless of territorial predisposition.
There are two explanations for why lab-reared males exhibited smaller medial and dorsal
cortices, compared to wild-caught males. One, lab-reared may lack normal cortical growth
during development. And two, cortical development proceeded normally, but eventually was
downregulated due to lack of environmental experiences. The fact that all lab-reared males had
similar cortical volume, regardless of territorial predisposition suggests that cortical volume may
be strongly regulated by experiential component, which is contrary to the authors' prediction.
The authors concluded that simplified environments, specifically those introduced in captivity,
significantly attenuates the medial and dorsal cortices in U ta stansburiana.
This is a novel experiment in that it is the first to investigate the combined roles of
territoriality and environment of the medial and dorsal cortices of lizards. The results of this
study are valuable for future research of the neuronal substrates of environmental as well as
phenotypic and genotypic influences. This study revealed that the size of this particular lizard
species are is directly proportional to the growth of both medial and dorsal cortices relative to
remainder of the telencephalon, regardless of territorial predisposition. Although, territorial
predisposition does have an effect when examining only size of cortices, in that the dorsal
cortex exhibited a larger size in the territorial wild-caught males, compared to the other three
groups (nonterritorial wild-caught males and territorial and nonterritorial laboratory reared
males).
One shortcoming of the experimental design in this study is that the territorial
phenotypes were not extremely similar in their territorial predisposition. Within the article, Table
1 indicates that between the territorial groups, the orange phenotype is highly aggressive and
the blue phenotype is somewhat territorial, although mostly in the context of mate guarding, and
that they are cooperative with other genotypes within their phenotype. The home range size of
blue phenotype is homologous to that of the nonterritorial yellow phenotype, at ~40m2, while the
territorial orange male has a home range size of ~100m2. This comparison is similar for
testosterone levels, with the orange phenotype exhibiting an average 40ng/ml higher than both
blue and yellow. It seems that the blue phenotype should not have been classified as
territorial, at least not at the same level as the orange phenotype. This fault in classification
may be the cause for territoriality not showing a significant effect on growth of the medial and
dorsal cortices.
Future studies would benefit from ensuring that the behavioral predispositions of their
subjects are adequately equal. Specifically, a future study could replicate this study, but with the
exception of analyzing the blue phenotype Uta stansburiana as a separate intermediate
territoriality. This would enable the experimenters to design a spectrum of neuronal substrate
response to territoriality and environmental experience. Building off these results will contribute
much needed information to understanding the effects of phenotype, genotype, and
environment on neuronal substrates in nonmammals.