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Vibrissa resonance as a transduction mechanism for tactile encoding.

We present evidence that resonance properties of rat vibrissae differentially amplify high-frequency and complex tactile signals. Consistent with a model of vibrissa mechanics, optical measurements of vibrissae revealed that their first mechanical resonance frequencies systematically varied from low (60-100 Hz) in longer, posterior vibrissae to high ( approximately 750 Hz) in shorter, anterior vibrissae. Resonance amplification of tactile input was observed in vivo and ex vivo, and in a variety of boundary conditions that are likely to occur during perception, including stimulation of the vibrissa with moving complex natural stimuli such as sandpaper. Vibrissae were underdamped, allowing for sharp tuning to resonance frequencies. Vibrissa resonance constitutes a potentially useful mechanism for perception of high-frequency and complex tactile signals. Amplification of small amplitude signals by resonance could facilitate detection of stimuli that would otherwise fail to drive neural activity. The systematic map of frequency sensitivity across the face could facilitate texture discrimination through somatotopic encoding of frequency content. These findings suggest strong parallels between vibrissa tactile processing and auditory encoding, in which the cochlea also uses resonance to amplify low-amplitude signals and to generate a spatial map of frequency sensitivity.

Animals↗

Unilateral stimulation or removal of rat vibrissae: analysis of nerve growth factor and tyrosine hydroxylase mRNA in the brain.

Previous work has shown that unilateral manipulation of vibrissae in the rat can lead to behavioral asymmetries and to neuronal changes in the basal ganglia: in brief, vibrissae stimulation led to increases in neostriatal dopamine release, whereas unilateral removal of vibrissae led to asymmetries in striatal afferents and to bilateral changes in mesencephalic dopamine mechanisms which were related to the occurrence of behavioral asymmetries and the later recovery therefrom. In the present study, the analysis of neuronal mechanisms possibly affected by vibrissae manipulation was extended to the nerve growth factor and the expression of tyrosine hydroxylase mRNA. Unilateral stimulation or removal of the vibrissae did not lead to significant changes in tissue levels of nerve growth factor in the neostriatum, parietal cortex (including the barrel cortex) or the hippocampus. In contrast, tyrosine hydroxylase mRNA in the substantia nigra and ventral tegmental area was affected by vibrissae removal but not by stimulation, as a bilateral increase in labeling was observed on the level of individual neurons. This effect was only observed in animals tested 4 h after vibrissae removal but not after 10 days. The results are discussed with respect to the interaction of vibrissae function with the basal ganglia, the neurotransmitter dopamine and mechanism of functional recovery.

Afferent Pathways↗

Mechanisms underlying experience-dependent potentiation and depression of vibrissae responses in barrel cortex.

Plasticity was studied in the barrel cortex of rats and mice. Vibrissae deprivation in adult and adolescent animals caused changes in the response properties of cortical neurons to stimulation of spared and deprived vibrissae. Plasticity involved both potentiation of spared vibrissae responses and depression of deprived vibrissae responses in layer II/III of the cortex. Vibrissae response potentiation was found to require alpha CaMKII in adult cortex. Vibrissae response depression was not found to occur in adult cortex but was found in adolescent animals for the principal vibrissa. Preliminary results suggested that vibrissae response depression exhibits both hetero- and homosynaptic components.

Animals↗

Coherent electrical activity between vibrissa sensory areas of cerebellum and neocortex is enhanced during free whisking.

We tested if coherent signaling between the sensory vibrissa areas of cerebellum and neocortex in rats was enhanced as they whisked in air. Whisking was accompanied by 5- to 15-Hz oscillations in the mystatial electromyogram, a measure of vibrissa position, and by 5- to 20-Hz oscillations in the differentially recorded local field potential (nablaLFP) within the vibrissa area of cerebellum and within the nablaLFP of primary sensory cortex. We observed that only 10% of the activity in either cerebellum or sensory neocortex was significantly phase-locked to rhythmic motion of the vibrissae; the extent of this modulation is in agreement with the results from previous single-unit measurements in sensory neocortex. In addition, we found that 40% of the activity in the vibrissa areas of cerebellum and neocortex was significantly coherent during periods of whisking. The relatively high level of coherence between these two brain areas, in comparison with their relatively low coherence with whisking per se, implies that the vibrissa areas of cerebellum and neocortex communicate in a manner that is incommensurate with whisking. To the extent that the vibrissa areas of cerebellum and neocortex communicate over the same frequency band as that used by whisking, these areas must multiplex electrical activity that is internal to the brain with activity that is that phase-locked to vibrissa sensory input.

Action Potentials↗

Vibrissa movement elicited by rhythmic electrical microstimulation to motor cortex in the aroused rat mimics exploratory whisking.

The rhythmic motor activity of the vibrissae that rodents use for the tactile localization of objects provides a model system for understanding patterned motor activity in mammals. Evidence suggests that neural circuitry in the brain stem provides rhythmic drive to the vibrissae. Yet multiple brain structures at higher levels of organization, including vibrissa primary motor cortex (M1), have direct projections to brain stem nuclei that are implicated in whisking. We thus asked whether output from M1 can control vibrissa movement on the approximately 10-Hz scale of the natural rhythmic movement of the vibrissae. Our assay of cortical control made use of periodic intracortical microstimulation (ICMS) to excite a region of vibrissa M1 cortex in awake, behaving animals and measurements of the stimulus-locked electromyogram (EMG) in both the intrinsic and extrinsic muscles that drive the vibrissae. We observed that ICMS evoked a prompt activation of the extrinsic muscles and a delayed and prolonged response in the intrinsic muscles. The relative timing and shape of these waveforms approximates the EMG waveforms seen during natural exploratory whisking. We further observed prompt activation of the intrinsic muscles, an occurrence not seen during exploratory whisking. Despite the latter difference in muscular activation, the motion of the vibrissae evoked by periodic ICMS strongly resembled the motion during natural, exploratory whisking. Interestingly, the extent of the movement was proportional to the level of arousal, as quantified by the amplitude of hippocampal activity in the theta frequency band. We interpret these data as demonstrating that M1 cortex can, in principle, initiate the full pattern of whisking on a cycle-by-cycle basis in aroused animals. Beyond issues of natural motor control, our result may bear on the design of algorithms for neuroprosthetic control of motor output.

Animals↗

Alterations in brainstem and cortical organization of rats sustaining prenatal vibrissa follicle lesions.

Vibrissa follicles were cauterized in late fetal or newborn rats to determine whether the relationships between brainstem and cortical changes observed after neonatal peripheral damage would also be obtained when vibrissa follicles were cauterized earlier in development. Vibrissa follicles were cauterized between embryonic day 15 (E-15) and the day of birth (P-0). The vibrissa-related representation in the brainstem was examined with cytochrome oxidase histochemistry, and that in the cortex was evaluated with either serotonin immunocytochemistry or anterograde labeling with Di-I when animals reached 6-8 days of age. There was a significant relationship between the ages at which lesions were carried out and the extent to which the representations of undamaged vibrissa follicles were altered in the brainstem and cortex. Peripheral lesions carried out between E-15 and E-18 resulted in significant increases in the cross-sectional areas of the patches corresponding to the undamaged vibrissa follicles in both the brainstem and cortex. Lesions at later ages resulted in significant increases only in the cortex. In some animals that sustained peripheral damage on E-20 and all of those that received lesions on P-0, there were aggregates of labeling in cortex that had no counterpart in the brainstem. Prenatal, but not postnatal, vibrissa follicle damage also reduced the overall dimensions of the cortical region devoted to the representation of these receptor organelles. Finally, there was a strong negative correlation between the magnitude of peripheral lesions (i.e., the number of vibrissa follicles ablated) and the extent of the reorganization in the brainstem and cortex.

Afferent Pathways↗

Differential fasciculation of follicular nerves for transferring specifically localized cues of the vibrissa rudiments to the central trigeminal sensory system in mice, as exploited with DiI and DiA labeling.

Vibrissa connections play a decisive role in setting the somatotopic coordinates in the trigeminal sensory system. Although previous studies have examined the development of peripheral patterning, certain questions are still in dispute, for example, the way vibrissa connections are structured, and the relationship between periphery and central organization, and ganglion cell organization. In order to fill the blanks left by previous studies, the extension of ganglionic branches and the formation of vibrissa connections were reexamined by using fluorescent carbocyanin dyes, DiI and DiA, during embryonic days 10 to 14 in mouse. Whole-mount preparations satisfactorily demonstrated the ganglionic fiber system, which allowed detailed analysis at both macroscopic and microscopic levels. We show here that the differential fasciculation of follicular nerves is the critical process for organizing vibrissa connections which modulate the initially extending fiber pattern. Follicular nerves developed by fine fibers arising from initially ordered root fascicles, so as to connect with vibrissa rudiments that developed on the facial prominences. During differential fasciculation, ganglion cells were segregated into distinct groups by the vibrissa connections, whereas central fiber terminals did not yet develop specific structures in the nuclear region. In the primary order of the trigeminal sensory system, vibrissa connections in the periphery were organized before those of the central structure. These results indicate that trigeminal ganglion cells have a critical binomial function in order to transfer the somatotopic relations among vibrissa rudiments into the topographic coordinates of the central system.

Animals↗

Somatotopic organization of vibrissae afferents in the trigeminal sensory nuclei of the rat studied by transganglionic transport of HRP.

Transganglionic transport of horseradish peroxidase (HRP) has been used to study the cell bodies and central projections of neurons innervating the vibrissae in the rat. These can be grouped into five horizontal rows and one posterior vertical row. Twenty-four to 48 hours after the nerves innervating different vibrissae were exposed to HRP, the trigeminal ganglia, brainstem, and upper cervical spinal cord were fixed by perfusion and serial sections were processed according to the tetramethylbenzidine technique. The results revealed a tendency for somatotopic organization in the trigeminal ganglion of cell bodies innervating the different vibrissae. Corresponding termination areas in the trigeminal sensory nuclei showed a detailed pattern of organization replicating the peripheral organization of the vibrissae. In all trigeminal sensory nuclei the horizontal rows are represented in an inverted fashion from dorsal to ventral, i.e., the most dorsal row is represented most ventrally. In addition, the more anterior a vibrissa is located, the deeper is it represented in the rostral nonlaminated nuclei. The situation is reversed in the laminated nucleus caudalis. The posterior vertical row is represented most superficially in the rostral nonlaminated nuclei, but most deeply in the laminated nucleus caudalis. In nucleus caudalis there are also rostrocaudal differences in the representation of different vibrissae. Thus, the posterior vibrissae in a horizontal row have their main representations more caudally than the anterior vibrissae. The posterior vertical row has its main representation most caudally, in the C1 segment.

Afferent Pathways↗

Vibrissae tactile stimulation: (14C) 2-deoxyglucose uptake in rat brainstem, thalamus, and cortex.

The right mystacial vibrissae of awake, adult rats were stroked at 4-6 times/second and brain regions which increased (14C) 2-deoxyglucose (2DG) uptake were mapped autoradiographically. The ventral parts of the ipsilateral spinal trigeminal nuclei pars caudalis (Sp5c), pars interpolaris (Sp5i), pars oralis (Sp5o), and the principal trigeminal sensory (Pr5) nuclei were activated. The lateral part of the ipsilateral facial (VII) nucleus (the region which innervates the vibrissae muscles) was also activated possibly via excitatory, trigeminal (Sp5c, Sp5i, Sp5o, Pr5) sensory afferents. A number of regions were activated contralateral to the sensory stimulus. Discrete patches of (14C) 2DG uptake occurred in deep layers of the superior colliculus (SCsgp). Dorsolateral and dorsomedial parts of the ventrobasal nucleus (VB), and posterior, dorsolateral parts of the reticular nucleus (R) of thalamus were activated, along with broad portions of the primary somatosensory cortex (SI) and second somatosensory cortex (SII). Though all layers of SI and SII cortex increased 2DG uptake, VB thalamic afferents to layers IV and Vc-Vla presumably accounted for the greater activation of these cortical layers during repetitive sensory stimulation of the vibrissae (RSSV). Activation of the above structures fits well with known anatomical data. However, the pattern of activation during RSSV was very different from that previously described during vibrissae motor cortex stimulation (VMIS). RSSV and VMIS both produced similar repetitive movements of all the mystacial vibrissae. However, only a few overlapping brain regions were activated during both RSSV and VMIS. These RSSV-VMIS overlap zones included Sp5o; rostral Sp5i; lateral VII; SCsgp; ventrobasal-posteromedial and ventrobasal-ventrolateral zones in thalamus; and a rostral region of SI probably anterior to the Woolsey vibrissae barrelfield in the dysgranular somatosensory (SI) cortex. Since RSSV and VMIS would both be expected to activate vibrissae proprioceptors, we have hypothesized that vibrissae proprioceptive input was processed in part in the RSSV-VMIS overlap zones. Convergence of motor-sensory inputs and other types of processing could have also occurred in these overlap zones.

Animals↗

Distributed representation of vibrissa movement in the upper layers of somatosensory cortex revealed with voltage-sensitive dyes.

We have identified large-scale patterns of electrical activity in circuits that occur in response to stimulation of peripheral receptors. Our focus was on primary (S1) vibrissal cortex of anesthetized rat, and we used optical techniques in conjunction with voltage-sensitive dyes to measure depolarization of the upper layers of cortex. Displacement of one vibrissa produced a field of activity that extends over very many cortical columns in S1. There are multiple, focal maxima within this field. A global maximum is located near the center of the field of activity, and, as determined electrically and histologically, this site maps to the cortical column appropriate for the deflected vibrissa. The amplitude of this component attains a steady-state value under continuous stimulation. Additional temporal characteristics are revealed by the response to a single displacement; the signal was triphasic and began with a prompt depolarization that was followed by a transient phase of inhibition and a final phase of long-lasting depolarization. The somatotopy of the other, satellite maxima in the field of activity were established through the reconstruction of the fields of activity produced by individual stimulation of other vibrissae. Local maxima for one vibrissa were seen to overlie the global maximum found for stimulation of nearest- and next-nearest-neighbor vibrissae. In contrast to the amplitude of the global maxima, the amplitude associated with the local maxima was not maintained with either continuous or infrequent but repetitive stimulation. Finally, the field of activity induced by alternate deflection of two neighboring vibrissae was suppressed in amplitude in comparison to the summed amplitudes of the signals elicited by deflection of each vibrissa alone. We suggest that these patterns of activity are a manifestation of the dynamic interaction among neighboring cortical columns.

Adaptation, Physiological↗

Reinnervation of a single vibrissa after nerve excision in the adult rat.

After a survival time of 180 days following the excision of a 2 mm segment of the vibrissal nerve to the gamma straddler vibrissa in the adult rat, a retrograde fluorescent single-labelling experiment revealed that 46% of the injured vibrissal sensory neurones had regenerated their peripheral processes. Peripheral collateral sprouting was not involved in the reinnervation of the denervated gamma vibrissa, as proved by a retrograde fluorescent double-labelling experiment. The regenerating nerve fibres did not invade the intact neighbouring vibrissae of the gamma vibrissa, and the sensory nerve fibres of the intact vibrissae were not translocated to the denervated gamma vibrissa. Thus, the sensory function of the denervated gamma vibrissa was restored exclusively by the regeneration of the damaged vibrissal nerve.

Animals↗

Curly vibrissae, a new mutation in the Wistar-derived rat.

A new mutation was identified in the PD (Preaxial Duplication) strain of rats, the main manifestations of which were curly and sparse vibrissae with retarded outer hair growth. As the main characteristic of this mutant rat is abnormally curled appearance of the vibrissae, "curly vibrissae" is proposed as the name of this mutant gene, and "cv" as its symbol. Genetic analyses reveal that the mutant characteristics are inherited as autosomal recessive traits and the cv gene is independent from the pd gene that carried by the original PD colony. The cv/cv homozygous rats have a small number of short and/or curly vibrissae around the nose. The vibrissae on the cheek and/or above the eyes are also short and curled; however, no vibrissa appears on the lower mandible. Although hair growth seems to be retarded, the outer hairs showed nearly normal length by 10 weeks of age. The outer hairs of matured cv/cv rats appear silky and translucent. The adult mutant rats often exhibit loss of hair on the head and/or back. Lactating females usually lose their abdominal hair. Both sexes of cv/cv homozygotes have normal reproductive ability. No internal malformations accompany vibrissa and hair abnormalities.

Alleles↗

Rhythmic whisking by rat: retraction as well as protraction of the vibrissae is under active muscular control.

The rhythmic motor activity of the vibrissae that rodents use for the tactile localization of objects provides a model system for understanding patterned motor activity in mammals. The muscles that drive this whisking are only partially fixed relative to bony attachments and thus shift their position along with the movement. As a means to characterize the pattern of muscular dynamics during different patterns of whisking, we recorded electromyogram (EMG) activity from the muscles that propel individual follicles, as well as EMG activity from a muscle group that moves the mystacial pad. The dominant pattern of whisking in our behavioral paradigm, referred to as exploratory whisking, consisted of large amplitude sweeps in the frequency range of 5-15 Hz. The frequency remained remarkably constant within a bout of whisking but changed values between bouts. The extrinsic musculature, which shifts the surface of the pad backwards, was found to be activated in approximate antiphase to that of the intrinsic muscles, which rotate individual vibrissae forward. Thus retraction of the vibrissae was driven by a backward shift in the attachment point of the follicles to the mystacial pad. In a less frequent pattern of whisking, referred to as foveal whisking, the vibrissae are thrust forward and palpate objects with low-amplitude movements that are in the higher frequency range of 15-25 Hz. Protraction of the vibrissae remains driven by the intrinsic muscles, while retraction in this pattern is largely passive. Interestingly, a mechanical argument suggests that activation of the extrinsic muscles during foveal whisking is not expected to affect the angle of the vibrissae. As a means to establish if the phasic control of the intrinsic versus extrinsic muscles depended on sensory feedback, we characterized whisking before and after bilateral transections of the infraorbital branch of the trigeminal sensory nerve. The loss of sensory feedback had no net effect on the antiphase relation between activation of the intrinsic versus extrinsic muscles over the full frequency range for exploratory whisking. These data point to the existence of a dual-phase central pattern generator that drives the vibrissae.

Animals↗

Central versus peripheral determinants of patterned spike activity in rat vibrissa cortex during whisking.

We report on the relationship between single-unit activity in primary somatosensory vibrissa cortex of rat and the rhythmic movement of vibrissae. Animals were trained to whisk freely in air in search of food. Electromyographic (EMG) recordings from the mystatial pads served as a reference for the position of the vibrissae. A fast, oscillatory component in single-unit spike trains is correlated with vibrissa position within the whisk cycle. The phase of the correlation for different units is broadly distributed. A second, slowly varying component of spike activity correlates with the amplitude of the whisk cycle. For some units, the phase and amplitude correlations were of sufficient strength to allow the position of the whiskers to be accurately predicted from a single spike train. To determine whether the observed patterned spike activity was driven by motion of the vibrissae, as opposed to central pathways, we reversibly blocked the contralateral facial motor nerve during the behavioral task so that the rat whisked only on the ipsilateral side. The ipsilateral EMG served as a reliable reference signal. The fast, oscillatory component of the spike-EMG correlation disappears when the facial motor nerve is blocked. This implies that the position of vibrissae within a cycle is encoded through direct sensory activation. The slowly varying component of the spike-EMG correlation is unaffected by the block. This implies that the amplitude of whisking is likely to be mediated by corollary discharge. Our results suggest that motor cortex does not relay a reference signal to sensory cortex for positional information of the vibrissae during whisking.

Action Potentials↗

Prevention of galanin upregulation following neonatal infraorbital nerve transection or attenuation of axoplasmic transport does not rescue central vibrissae-related patterns in the rat.

Neonatal transection of, or blockade of axoplasmic transport in, the infraorbital nerve [ION, the trigeminal (V) branch that supplies the mystacial vibrissae follicles] results in a loss of all central patterns corresponding to the vibrissae follicles in the brainstem, thalamus and cortex except for those of the central terminal arbors of ION primary afferents that survive this lesion. Both of these manipulations also result in a rapid and dramatic upregulation of at least two peptides, galanin and neuropeptide Y, in surviving vibrissae-related primary afferents. Galanin is of particular interest, because this peptide has effects on neuronal activity and growth, both factors which may be involved in the disappearance of central vibrissae-related patterns in rats that have sustained neonatal ION transection or axoplasmic transport blockade. The present study used antisense technology to determine whether the upregulation of galanin in the central terminals of ION primary afferents is necessary for the loss of central vibrissae-related patterns in rats. Newborn rats had their left ION transected or axoplasmic transport in this nerve blocked by application of a vinblastine-impregnated implant, and at the same time received an injection of commercially synthesized phosphorothioate oligodeoxynucleotide sequences (15-20 bases) directly into the V ganglion in order to block galanin upregulation. These injections effectively prevented the upregulation of this peptide which is normally associated with ION transection or axoplasmic transport blockade. Preventing galanin upregulation, however, did not prevent or attenuate the loss of central vibrissae-related patterns in the brainstem or cortex normally observed following ION transection or axoplasmic transport blockade in this nerve. These results are thus consistent with the conclusion that the upregulation of galanin in the central terminals of V primary afferents, observed after damage to or attenuation of axoplasmic transport in the ION, is not necessary for the reorganization that results in a disappearance of central vibrissae-related patterns in the V neuraxis.

Animals↗

Vibrissae representation in subcortical trigeminal centers of the neonatal rat.

In the neonatal rat differential activity levels of the metabolic enzyme succinic dehydrogenase (SDH) reveal intricately detailed sgementation in the neuropil of the spinal and principal trigeminal nuclei of the brainstem and in the ventrobasal complex of the thalamus. The segmentation occurs in the portions of these nuclei that electrophysiological evidence has indicated to be related to the mystacial vibrissae and sinus hairs on the face of the rat. Indeed, the pattern of segmentation in each nucleus replicates the topographic distribution of the vibrissae and sinus hairs. Further, within the spinal trigeminal nucleus, there appear to be two distinct representations of the vibrissae, one in the subnucleus caudalis and a second in the subnucleus interpolaris. Examination of these patterns of segmentation indicates that the large mystacial vibriaase and sinus hairs on the face of the young rat are somatotopically represented three times within the trigeminal complex, as straight cylinders of neuropil, and once in the ventrobasal complex, as curved cylinders of neuropil. Neonatal vibrissae damage leads to an aberrant organization of the segmentation in the spinal trigeminal nucleus and the ventrobasal complex. In the spinal trigeminal nucleus, the SDH activity in areas associated with damaged vibrissae is of a lower than normal density, and patterns are indistinct. However, rows of clusters associated with the adjacent normal vibrissae are apparent and appear to be enlarged. In the ventrobasal complex, vibrissae damage results in bands of normal density SDH activity where rows of segmented clusters would normally be present. Comparison of these data to the cortical data in the previous paper (Killackey and Belford, '79) indicates that cortical and nuclear structures can have aspects of their development controlled by similar mechanisms.

Animals↗

The development of vibrissae representation in subcortical trigeminal centers of the neonatal rat.

In every station of the trigeminal system of the young rat, the segmented activity of the mitochondrial enzyme succinic dehydrogenase (SDH) clearly delineates the representation of the mystacial vibrissae. In the trigeminal complex of the medulla, three parallel representation can be seen, two in the spinal trigeminal nucleus and one in the principal trigeminal nucleus. In the next station, the ventrobasal complex of the thalamus, a single representation occurs. Likewise, layer IV of somatosensory cortex contains one representation of the vibrissae. Further, neonatal damage to the mystacial vibrissae results in anomalies within each representation. The present study delineates both the normal development of subcortical trigeminal stations and the aberrant organization seen after vibrisse removal. The results of a similar study on somatosensory cortex (Killackey and Belford, '79) and the present data allow the comparison of the development of each of the five vibrissae representations in the trigeminal system. In the brainstem, each of the three trigeminal complex representations are present at birth, although the pattern becomes more distinct over the first several days of life. Interestingly, vibrissae removal at birth induces an aberrant pattern that is distinct by postnatal Day 3. Although details are not equally discernible in each representation, the abnormalities appear to be similar. The SDH segmentation in the ventrobasal complex develops during postnatal Days 1 through 4. At Day 1, portions of the matrix of high density SDH activity break up into bands. Clusters can be discerned within these bands on Day 2. By Day 4 the pattern is sharply delineated. Vibrissae removal at birth results in anomalies that are a part of the initial development of segmentation, not a later reorganization. Comparison of the present data with that of our previous studies indicates that there is a sequential development of the central somatosensory structures related to the vibrissae, beginning with the most peripheral station. Further, there are many similarities in the development of each station. There are also differences which are particularly important in comparing the trigeminal nuclei with the later stations. The unique features in the abnormal development of the trigeminal nuclei are likely due to their direct connections with the periphery.

Animals↗

Effect of castration and testosterone propionate on mouse vibrissae.

Vibrissae of normal mice were measured for six cycles starting at birth. Groups of immature male albino mice were castrated at 21 days of age when E, F, G and H (Danforth nomenclature, 1925) vibrissae were in their first cycle of growth. Levels of testosterone in their peripheral plasma were reduced to about one tenth of the normal. The final length achieved by their vibrissae in all subsequent cycles was shorter than that of normal mice. Another group of immature male mice were treated with implants of testosterone propionate to give plasma levels of approximately five to ten times normal. Their vibrissae showed a significant increase in final length in all growth cycles before and after sexual maturation. Vibrissae of adult male mice treated with excess testosterone showed a significant increase in final length at the end of the growth cycle immediately after commencement of treatment. On the other hand, vibrissae of castrated adult mice showed a significant decrease in final length in the second successive cycle after castration. The rate of growth of all vibrissae remained unchanged. Changes in their final length were brought about by changes in the period of growth. Castrated mice had a slightly longer resting period.

Animals↗