Neuroactive substances in the developing dorsomedial telencephalon of the pigeon (Columba livia): differential distribution and time course of maturation

J Comp Neurol. 1994 Jul 22;345(4):537-61. doi: 10.1002/cne.903450406.


The avian hippocampal formation has previously been shown to contain many of the same neurotransmitters and related enzymes that are found in mammals. In order to determine whether the relatively delayed development of the mammalian hippocampus is typical of other vertebrates, we investigated the maturation of a variety of neuroactive substances in the hippocampal formation of the homing pigeon. The distribution of two transmitter-related enzymes, choline acetyltransferase (ChAT) and tyrosine hydroxylase (TH), the neurotransmitter GABA, and four neuropeptides (substance P, enkephalin, neuropeptide Y, and somatostatin) was studied by immunohistochemistry in the developing hippocampal complex. The pattern and/or the time course of changes in the distribution of immunoreactivity varied among the different neuroactive substances examined. Immunoreactivity to ChAT and TH was found exclusively in fibers and terminal-like processes, whereas GABA and peptide immunoreactivity was seen in cells and neuropil. Quantitative differences in the density, number, and size of stained cells were assessed by a computer-assisted image analyzer. For the majority of the substances, developmental patterns in the distribution of immunoreactivity differ between the hippocampus proper and the area parahippocampalis, the two major areas that together make up the avian hippocampal complex. The adult pattern of immunoreactivity was generally attained by 3 weeks after hatching. For many of the neuroactive substances found in cell bodies, there was a gradual decrease in the density of immunoreactive cells with a concomitant increase in the density of immunoreactive neuropil. The actual number of stained cells usually increased to a peak at 9 days posthatching and then declined until 3 weeks posthatching, when the adult value was reached. These results are discussed in relation to the advantages that the pigeon hippocampal complex may provide in the study of developmental processes. Parallels with the distribution of the same neuroactive substances in the mammalian hippocampus are used to suggest possible functional similarities between the avian and mammalian hippocampal regions.

Publication types

  • Comparative Study
  • Research Support, Non-U.S. Gov't
  • Research Support, U.S. Gov't, P.H.S.

MeSH terms

  • Animals
  • Choline O-Acetyltransferase / analysis*
  • Columbidae / metabolism*
  • Enkephalins / analysis
  • Hippocampus / chemistry*
  • Hippocampus / growth & development
  • Hippocampus / metabolism
  • Immunohistochemistry
  • Neuropeptide Y / analysis
  • Neuropeptides / analysis*
  • Neurotransmitter Agents / analysis*
  • Somatostatin / analysis
  • Substance P / analysis
  • Time Factors
  • Tyrosine 3-Monooxygenase / analysis*
  • gamma-Aminobutyric Acid / analysis


  • Enkephalins
  • Neuropeptide Y
  • Neuropeptides
  • Neurotransmitter Agents
  • Substance P
  • Somatostatin
  • gamma-Aminobutyric Acid
  • Tyrosine 3-Monooxygenase
  • Choline O-Acetyltransferase