The best understood functional activity of the pineal gland is its diurnal production of melatonin in response to environmental lighting cues. Several enzymes of the melatonin pathway respond to daily photoperiod changes, for example hydroxyindole-O-methyltransferase (HIOMT) and serotonin N-acetyltransferase (SNAT). Increased levels of the glycolytic enzyme neuron-specific enolase (NSE) are thought to reflect increased physiological demands placed on neurons and neuroendocrine tissues. Homodimer non-neuronal enolase isoenzyme (NNE) is immunolocalized to cells, and the hybrid enolase (consisting of subunits from NSE and NNE) has been seen in cerebellar stellate and basket cells. Although not rate limiting, concentrations of both NSE and NNE in the rat pineal gland rival that of the forebrain. The rat pinealocytes demonstrates a mosaic pattern of immunostaining for the NSE antigen. Furthermore, an increasing proximal to distal immunostaining gradient has been reported in the gland. In this study, it was hypothesized that structural and immunohistochemical differences represent pinealocyte heterogenous functional activity. This was tested in two rodent populations (rats and Djungarian hamsters) by means of evaluating the effect of photoperiod manipulation on morphological development and immunohistochemical localization of enolase isoenzymes. Djungarian hamsters respond well to photoperiod changes, while laboratory rats are thought to be insensitive to environmental lighting changes. The effect of photoperiod in the hamster was confirmed by significantly different (p < 0.01) body and testes weights for animals housed under the two different photoperiods (long day, 16L:8D; short day 6L:18D). In addition, rats exposed to the long day photoperiod (12L:12D) had significantly greater body and testes weights (p < 0.05) than animals raised under the short day photoperiod (6L:18D). Photoperiod also influenced pineal gland soluble protein, protein concentration, and NSE concentration in both species. Generally, the levels were greatest in extracts from the glands of animals subjected to the short day environment. Gel electrophoresis revealed a protein or group of proteins that were particularly responsive to changes in environmental lighting; the estimated molecular weight was 80 kD. In both species, NNE exhibited the same heterogenous immunostaining as NSE, and was localized to the same pinealocytes in adjacent sections. The preceding strongly suggests that the hybrid enolase is found in the rat and hamster pineal gland. If so, this may reflect a balance between simultaneous expression of the NSE and NNE genes. The NSE increasing proximal to distal immunostaining gradient was repeated by NNE in the rat gland, but it was in the opposite direction for the hamster. Morphological features, however, may have contributed to the gradient nature of enolase isoenzyme immunostaining, in that regional nuclear and blood vessel volume fractions were influenced by photoperiod.

LLU Discipline





Graduate School

First Advisor

Paul J. McMillan

Second Advisor

Robert L. Schultz

Third Advisor

Paul C. Engen

Fourth Advisor

Benjamin H.S. Lau

Fifth Advisor

Steven M. Yellon

Degree Name

Doctor of Philosophy (PhD)

Degree Level


Year Degree Awarded


Date (Title Page)




Library of Congress/MESH Subject Headings

Pineal Body -- physiology; Lighting; Phosphopyruvate Hydratase -- physiology



Page Count

xii; 158

Digital Format


Digital Publisher

Loma Linda University Libraries

Usage Rights

This title appears here courtesy of the author, who has granted Loma Linda University a limited, non-exclusive right to make this publication available to the public. The author retains all other copyrights.


Loma Linda University Electronic Theses and Dissertations

Collection Website



Loma Linda University. Del E. Webb Memorial Library. University Archives