Biomedical subjects
Publications and source records attributed to D W Pfaff.
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Inputs to medial longissimus (ML) and lateral longissimus (LL) motoneurons were studied in urethane or urethane-chloralose anesthetized rats by recording from ML and LL nerves while stimulating ipsilateral lumbosacral dorsal roots, medial medullary reticular formation (RF), vestibular nuclei (VN), dorsal midbrain (MDBR), or ventromedial hypothalamus (VMH). Stimulation of appropriate dorsal roots produced short-latency responses (1.5-3.0 ms) in nerves to medial longissimus or lateral longissimus. The connections underlying these responses, which could be monosynaptic, are weak, since generally two or more stimuli were necessary for a response to occur. Short-latency LL nerve responses required more dorsal root stimuli than did ML nerve responses and stable LL responses sometimes could not be obtained, suggesting that segmental reflexes to a back muscle (LL) could be weaker than those to a proximal tail muscle (ML). Trains of conditioning stimuli delivered to the RF, VN, and MDBR facilitated segmental responses in ML nerves or LL nerves. Temporal profiles of facilitation of ML differed for the three regions. On one extreme, the facilitation produced by RF conditioning required few stimuli (median, 3 shocks) and peak facilitation occurred at short condition-test intervals (median, 1.5 ms). On the other extreme, facilitation produced by MDBR conditioning required long trains (median, 14 stimuli) and peak facilitation occurred at longer condition-test intervals (median, 10 ms). Stimulation within the VMH never facilitated ML or LL nerve activity. These results demonstrate excitatory connections from reticular formation, vestibular nuclei and the dorsal midbrain to medial longissimus and lateral longissimus. Such connections could be involved in behaviors mediated by midbrain, and in postural regulation through brain stem control of axial musculature. Motoneuron cell bodies for LL, ML and lumbar transversospinalis (TS) muscles were localized by ejecting dye at sites where unitary antidromic responses to muscle nerve stimulation were recorded extracellularly. ML cells were found ventrolaterally in the L6-S1 ventral horn. LL and TS cells were found medially in the ventral horn of the lumbar enlargement.
Neurons in the hypothalamus, other diencephalic regions, and the telencephalon which project to the mesencephalic central gray (CG) and the region lateral to it were demonstrated, in the rat, by the horseradish peroxidase retrograde neuroanatomical tracing method with diaminobenzidine and tetramethyl benzidine visualization reactions. The greatest concentrations of neurons that project to the dorsal mesencephalon were found in the ventromedial nucleus, particularly the anterior and ventrolateral subdivisions, in the dorsal premammillary nucleus, and in the zona incerta. Neurons that project to or lateral to the CG were also found in the laterocaudal hypothalamus, the dorsomedial hypothalamus, regions of the anterior hypothalamic area, specific areas of the cerebral cortex (32, 29, 8, 8A, 13, 14), and the central nucleus of the amygdala. Some neurons that project were also found in the preoptic area, septum, bed nucleus of the stria terminals, and the habenula. More neurons in the mediocaudal quadrant of the hypothalamus project to the mesencephalon than do those in laterocaudal, mediorostral, or laterorostral quadrants. More neurons in the medial than the lateral half, and more in the caudal than the rostral half of the hypothalamus project to the mesencephalon. More neurons project to the central gray, or the region lateral to it, at the levels of the superior colliculus, or intercollicular region, than at the level of the inferior colliculus. These descending connections to the midbrain, particularly from the hypothalamus and zona incerta, are probably components of neural networks that regulate nociception, certain neuroendocrine functions, sexual and other behaviors, and certain autonomic functions.
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A cell-by-cell analysis of the magnocellular elements in hypothalami of fifty Long-Evans (normal) and Brattleboro (diabetes insipidus) rats was done using the unlabeled antibody enzyme technique (PAP) with primary antisera directed against oxytocin (OXY), vasopressin (ADH), and the neurophysins. The magnocellular neurons of the hypothalamus were found in the supraoptic (SON), paraventricular (PVN), and anterior commissural (ACN) nuclei, a number of accessory nuclei, and as individual cells in the anterior hypothalamic area. SON was divided by the optic tract into the principal part and retrochiasmatic SON. In retrochiasmatic SON a majority of the cells contained vasopressin. Within the principal part of SON oxytocin-producing cells tended to be found rostrally and dorsally, while the vasopressin cells were more common caudally and ventrally. PVN was divided into three subnuclei, the medial, lateral, and posterior subnuclei, on the basis of cellular morphology and peptide content. The magnocellular cells of the medial and lateral PVN were closely packed together and nearly round, while those of posterior PVN were more separated and fusiform in shape with their long axis running in a medio-lateral direction. Medial PVN consisted primarily of oxytocin-producing cells, while lateral PVN was formed by a core of vasopressin-producing cells with a rim of oxytocin cells. Posterior PVN contained largely oxytocin-producing cells. Both ADH and OXY cells were found in the accessory nuclei. In the Long-Evans rat the SON had, on the average, 1443 OXY and 3236 ADH cells; the PVN had 1174 OXY and 976 ADH cells; and the accessory magnocellular groups in the hypothalamus (including the ACN) had 1286 OXY and 552 ADH cells. The Brattleboro strain animal had similar numbers of cells in these nuclei. (The cells which contain ADH in normal animals were identified in the Brattleboro rat as large, neurophysin-negative cells.) Thus, a large fraction of the magnocellular oxytocin- and vasopressin-producing cells in the rat were located outside of the PVN and SON. One accessory cell group in particular, ACN, had 616 OXY cells, or about 50% as many as PVN. In each nucleus the sum of the numbers of OXY and ADH cells was approximately the number of neurophysin cells.
The simplest mechanism for lordosis behavior, as allowed by the present data, is as follows. Estrogen acts on the ventromedial nucleus of hypothalamus (VMN) to induce a tonic facilitatory output, which is relayed by neurons in mesencephalic central gray and then the medullary reticulospinal tract to the spinal cord. To trigger the lordosis reflex, sensory inputs from cutaneous touch-pressure receptors located in rump-perineal skin are required. Neural pathways exist for transmitting these inputs to motor neurons in the spinal cord and to certain supraspinal locations, but our electrophysiological studies indicate that the most likely place where these sensory inputs interact with estrogen-influenced neural activity to elicit the reflex is the spinal cord itself.
The fine structure of the ventrolateral and dorsomedial subdivisions of the ventromedial nucleus (VMN) of the hypothalamus was examined in ovariectomized/control and ovariectomized/estrogen-treated rats to compare neurons of these areas to other neurons (specifically the ventrolateral thalamus), and to determine the effects of estrogen on these cells. The neurons of the VMN contain a large nucleus with a prominent nucleolus, rough endoplasmic reticulum (RER), polysomes, a Golgi complex, coated, uncoated and dense-cored vesicles, lysosome-like bodies, inclusion bodies, multivesicular bodies whorl bodies and myelin figures. Similar organelles were present in the neurons of the ventrolateral thalamus, although polysomes wee more prominent, and the cells lacked dense-cored vesicles in the perikarya. Differences in the cells of the VMN between ovariectomized/control and ovariectomized/estrogen-treated rats included a more conspicuous stacking of the RER and greater number of dense-cored vesicles in the estrogen-treated group in both the ventrolateral and dorsomedial subdivisions. In both areas the differences were statistically significant, although more marked in the ventrolateral subdivision. In both VMN subdivisions, the increased stacking of the RER could be correlated with the greater number of dense-cored vesicles and may reflect increased biosynthesis of a secretory product.
The effect of water deprivation or estrogen treatment on the oxytocin content of rat hypothalamic cells was examined using a quantitative immunohistological technique. Oxytocin-containing cells were visualized using the immunoperoxidase technique of Sternberg and a primary antiserum directed against oxytocin. The optical density of the darkest 3.2 micrometer diameter spot in the cytoplasm of a cell was used as a measure of the oxytocin content of that cell. Water deprivation produced a significant decrease in anti-oxytocin staining in the anterior commissural nucleus of males and females. There was a similar decrease in the paraventricular nucleus of males, but not in the paraventricular nucleus of females or the supraoptic nucleus of either males or females. Estrogen treatment of ovariectomized female rats produced a fall in anti-oxytocin staining in the anterior commissural, but not paraventricular or supraoptic nuclei.
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The distribution of estrogen-concentrating cells in the rat brain has been previously described in detail as has the distribution of cells containing oxytocin, vasopressin, and their associated neurophysins. This report describes the simultaneous localization of [3H]estradiol by steroid autoradiography and neurophysin by immunohistology in cells in the rat hypothalamus. In the rat, neurophysin-containing cells were found in three major nuclei - the supraoptic nucleus, the paraventricular nucleus (PVN) and the anterior commissural nucleus - as well as in a number of accessory nuclei and as individual cells in the anterior hypothalamic area. Of these, only the PVN has appreciable numbers of cells which both concentrate estradiol and contain neurophysin. Within the PVN these cells are found predominantly in the posterior subnucleus.
Four methods used for the neurohistochemical demonstration of horseradish peroxidase (HRP) were quantitatively compared by counting retrogradely labeled neurons found after each method was used. HRP used as a retrograde marker is an important neuroanatomical tracing method, and maximum sensitivity in its demonstration of retrogradely, labeled neurons is important if these neuroanatomical studies are to completely demonstrate afferent neurons. The four methods compared were a diaminobenzidine (DAB) procedure, a Hanker-Yates procedure using P-phenylenediamine and pyrocatechol, an o-dianisidine procedure, and a tetramethyl benzidine (TMB) procedure. The TMB procedure resulted in a more complete topography of neurons afferent to the HRP application site, and demonstrated many more neurons in all afferent cell groups that either of the three other procedures. Use of the TMB method was especially critical in the cases of small HRP applications, a size useful for neuroanatomical studies, where the other methods demonstrated very few or no retrogradely labeled neurons. Neurons were judged to be retrogradely HRP labeled if they had small granules of the reaction product (the color varying with the chromogen) describing the somal shape, usually extending into the processes, and a clear nucleus. In addition, after the o-dianisidine or the TMB reaction a small number of retrogradely labeled neurons had soma and processes especially well filled with reaction product, giving the appearance of neurons from Golgi preparations. For a sensitive TMB reaction giving good results, exact H2O2 concentration, freshly prepared solutions, minimal postreaction exposure to alcohol, counterstaining, and clean glassware were each found to be important.
The lordosis reflex, which characterises female reproductive behaviour in rodents, can be potentiated in oestrogen-primed ovariectomised female rats by electrical stimulation of the ventromedial nucleus of the hypothalamus (VMN). Subsequent demonstration of an immediate and large facilitation of the lordosis reflex from the mesencephalic central grey (CG) prompted us to speculate that the VMN may exert an oestrogen-dependent tonic bias on the mesencephalic circuitry for lordosis via its heavy descending projections. Moreover, we considered that luteinising hormone-releasing hormone (LH-RH) could be involved in this system. LH-RH can potentiate lordosis when given systemically to oestrogen-primed ovariectomised rats, and neurotropic effects of this peptide have been shown by microiontophoresis. Also, some axons in the CG have been stained with antiserum to LH-RH. The present study examined a possible role of LH-RH in the CG in the regulation of the lordosis reflex of oestrogen-primed ovariectomised female rats. Infusion of exogenous LH-RH in the CG had an immediate facilitative effect on the lordosis reflex, whereas passive immunisation against endogenous LH-RH by anti-LH-RH gamma-globulin diminished the reflex.
The distribution of labeled cells in the chaffinch brain has been examined by autoradiography after an intramuscular injection of [3H]testosterone. In the telencephalon labeled cells were particularly concentrated in nucleus magnocellularis neostriatalis anterioris, nucleus septalis lateralis, and hyperstriatum ventrale, pars caudale. In the diencephalon the nucleus periventricularis magnocellularis, the nucleus magnocellularis posterioris and the area infundibularis contained many heavily labeled cells. In the mesencephalon and medulla, labeled cells were found in the nucleus intercollicularis and in the nucleus nervi hypoglossi, respectively. Measurements of the density of grains per cell revealed (1) clear quantitative differences in the extent of labeling in different regions of the brain and (2) a wide variation in the density of labeling of individual cells within a region. The relationship between these sites of hormone binding and known sites of androgen action in the avian brain is discussed.
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Electrical stimulation (ES) through fine-wire electrodes chronically implanted in the preoptic area and anterior hypothalamus suppressed the lordosis response in freely moving hormone-primed female hamsters. ES at these sites never induced or facilitated lordosis. When behavioral testing was completed small lesions were made at the tips of the electrodes, and the Fink-Heimer method was used to trace degenerating axons away from the sites of ES. This was done in an effort to determine the neural pathways important for the modulation of this behavior. The hypothalamic distribution of the stria terminalis was also charted for the hamster and was found to be similar to that in the rat, including projections to both the shell and core of the ventromedial nucleus (VMN). No single pattern of degeneration was common to all females showing ES-produced lordosis suppression. These data demonstrate that stimulation of several different pathways can produce behavioral states that are incompatible with the lordosis response. We suggest that ES in the medial preoptic-medial anterior hypothalamic continuum suppresses sexual receptivity by influencing neurons in the VMN region, while ES through more laterally placed electrodes suppresses receptivity by producing an incompatible state of behavioral activation, without directly influencing VMN region neural activity.