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PubMed1992

Difference in receptive field features of taste neurons in rat granular and dysgranular insular cortices.

Receptive fields (RFs) of 59 cortical taste neurons (35 in the granular insular area, area GI, 21 in the dysgranular insular area, area DI, and 3 in the agranular insular area, area AI) were identified in the oral cavity of the rat. The fraction of the neurons with RFs in the anterior oral cavity only was significantly larger in area GI (74.3%) than in area DI (42.9%). On the other hand, the fraction of neurons with RFs in both the anterior and posterior oral cavity was larger in area DI (42.9%) than in area GI (11.4%). On the whole, it is suggested that area GI is involved in discrimination of several taste stimuli in the oral cavity, whereas in area DI taste information originating from various regions of the oral cavity is integrated. When neurons were classified according to the best stimulus which most excited the neuron among the four basic tastes, different categories of taste neurons had RFs in different parts of the oral cavity. It is suggested that, in either taste area, different categories of taste neurons are involved in different sorts of taste coding. The majority of neurons in both areas had bilateral RFs. In area GI, neurons with RFs on single subpopulations of taste buds were significantly more numerous at the rostral region of the cortex than at the caudal region. There was no such relation between RF types and cortical localization in area DI. Otherwise, topographic representation of the oral cavity by taste neurons on the cortical surface was not obvious. RF features of taste neurons did not differ across layers in either cortical area.

Animals↗
PubMed1985

Responses of solitary tract nucleus neurons to taste and mechanical stimulations of the oral cavity in decerebrate rats.

Physiological characteristics of 45 taste and 15 mechanoreceptive units were examined in the solitary tract nucleus (NTS) of rats decerebrated at the pre- or midcollicular level, and compared with previous findings in the intact rat. The rostro-caudal extent of the area, where taste and mechanoreceptive neurons were recorded, was almost the same in the decerebrate rat as that in intact rat. The spontaneous discharge rate was significantly lower in the decerebrate rat than in the intact rat. The taste profile of the NTS units in decerebrate rats was quite different from that in intact rats; significant decreases in correlation coefficients were found between certain pairs of taste stimuli and spontaneous discharge rate, e.g. NaCl-quinine, sucrose-quinine. A large number of taste (18 of 31) and mechanoreceptive (12 of 15) units examined had receptive fields (RFs) on the palate, and four taste and two mechanoreceptive units on the circumvallate area. This contrasts with the findings in the intact rat. Some taste (n = 1) and mechanoreceptive units (n = 2) had large RFs. Taste units with different RF locations showed different taste profiles. Acute i.v. injection of amobarbital sodium affected only the response magnitude of taste units, suggesting that most of the differences between intact and decerebrate rats might be caused by decerebration. The present findings indicate that neural structures above the pre- or midcollicular level have tonic inhibitory or facilitatory effects on the response properties of NTS taste units.

Amobarbital↗
PubMed2006

Effects of acetylcholine on coding of taste information in the primary gustatory cortex in rats.

Acetylcholine (ACh) receptors are widely distributed throughout the cerebral cortex in rats. Recently, cholinergic innervation of the gustatory cortex (GC) was reported to be involved in certain taste learning in rats. Here, the effects of iontophoretic application of ACh on the response properties of GC neurons were studied in urethane-anesthetized rats. ACh affected spontaneous discharges in a small fraction of taste neurons (11 of 86 neurons tested), but influenced taste responses in 27 of 43 neurons tested. No correlations with ACh susceptibility were noted for spontaneous discharges and taste responses. Among the 27 neurons, ACh facilitated taste responses in 13, inhibited taste responses in 13 and either facilitated or inhibited taste responses depending on the stimuli in 1. Furthermore, ACh affected the responses to best stimuli that produced the largest responses among four basic tastants (best responses) in 7 of 27 taste neurons, to non-best responses in 9, and to both best and non-best responses in 11. ACh mostly inhibited the best responses (13 of 18 neurons). Thus, ACh often decreased the response selectivity to the four basic tastants and changed the response profile. Atropine, a general antagonist of muscarinic receptors, antagonized ACh actions on taste responses or displayed the opposite effects on taste responses to ACh actions in two-thirds of the neurons tested. These findings indicate that ACh mostly modulates taste responses through muscarinic receptors, and suggest that ACh shifts the state of the neuron network in the GC, in terms of the response selectivities and response profiles.

Acetylcholine↗
PubMed1982

Studies on the perception of taste: do primaries exist?

The idea that there are four primary tastes is basic to almost all research in gustation. Although this concept guides the formulation and interpretation of studies of taste psychophysics, as well as studies of taste stimuli, receptors, and neural organization, there is a surprising absence of studies designed to determine if this quadripartite view of taste is correct. The present group of three studies is designed to ask this question on a psychophysical level, comparing the taste data with combinations of tones, which appear to remain separate; subjects are asked: (a) whether tastes and tones--including mixtures--are perceived as singular or more-than-one, (b) whether increasing the number of components in a mixture (tastes or tones) results in increases in the perceived complexity, and (c) whether stimuli (tastes or tones) remain identifiable in mixtures with other stimuli. In all three experiments, it appears that the individual tones remained distinct and unaltered in combinations. Taste stimuli combined very differently from tones, in ways that suggested that the original components lost their identity, and sensations other than those in the original components (e.g., "primary tastes") were synthesized. From this it follows that taste is composed of many sensations, not just the "primary four."

Adolescent↗
PubMedDate not supplied

The neural code for taste in the brain stem: response profiles.

In the study of the neural code for taste, two theories have dominated the literature: the across neuron pattern (ANP), and the labeled line theories. Both of these theories are based on the observations that taste cells are multisensitive across a variety of different taste stimuli. Given a fixed array of taste stimuli, a cell's particular set of sensitivities defines its response profile. The characteristics of response profiles are the basis of both major theories of coding. In reviewing the literature, it is apparent that response profiles are an expression of a complex interplay of excitatory and inhibitory inputs that derive from cells with a wide variety of sensitivity patterns. These observations suggest that, in the absence of inhibition, taste cells might be potentially responsive to all taste stimuli. Several studies also suggest that response profiles can be influenced by the taste context, defined as the taste stimulus presented just before or simultaneously with another, under which they are recorded. A theory, called dynamic coding, was proposed to account for context dependency of taste response profiles. In this theory, those cells that are unaffected by taste context would provide the signal, i.e., the information-containing portion of the ANP, and those cells whose responses are context dependent would provide noise, i.e., less stimulus specific information. When singular taste stimuli are presented, noise cells would provide amplification of the signal, and when complex mixtures are presented, the responses of the noise cells would be suppressed (depending on the particular combination of tastants), and the ratio of signal to noise would be enhanced.

Animals↗
PubMed1987

What kind of exposure reduces children's food neophobia? Looking vs. tasting.

To determine the relative effectiveness of two different types of exposure on young children's preference for initially novel foods, 51 two- to five-year-old children received either "look" or "taste" exposures to seven novel fruits. Foods were exposed five, 10 or 15 times, and one food remained novel. Following the exposures, children made two judgments of each of the 21 pairs: one based on looking, the other on tasting the foods. Thurstone Case V scaling solutions were correlated with exposure frequency, and these were significant for the visual judgments of the looked at foods (r = 0.91), the visual judgments of the tasted (and looked at) foods (r = 0.97) and the taste judgments of the tasted foods (r = 0.94). The only non-significant relationship was for the taste judgments of the looked at (but never tasted) foods (r = 0.24). The results indicate that to obtain significant positive changes in preference, experience with the food must include experience in the modality that is relevant for the judgments. While visual experience produced enhanced visual preference judgments, visual experience was not sufficient to produce significantly enhanced taste preferences. This finding is consistent with a "learned safety" interpretation of the exposure effects noted in the taste judgments: experiences with novel tastes that are not followed by negative gastrointestinal consequences can produce enhanced taste preference.

Child, Preschool↗
PubMed2001

Role of parabrachial nucleus in submandibular salivary secretion induced by bitter taste stimulation in rats.

When rats lick a bitter taste solution such as quinine-hydrochloride, they secrete profuse amounts of saliva. The salivation has a higher flow rate than that induced by other qualities of taste stimulation: sweet, salty, and sour. The present study is aimed to clarify the neural mechanism of the quinine-evoked salivation by means of behavioral, neuroanatomical, and electrophysiological experiments. Behaviorally, submandibular salivary secretion and rejection behavior (gaping) were observed in normal rats, as well as in rats chronically decerebrated at the precollicular level. In chronically decerebrate rats, these quinine-evoked reactions were strongly suppressed by destruction of the medial part of the parabrachial nucleus, including the so-called taste area, and ventral part of the parabrachial nucleus, including the pontine reticular formation. Neuroanatomical study using a retrograde tracer, Fluoro-gold, revealed that the neurons sending their axons to the superior salivatory nucleus, parasympathetic secretory center, were located mainly in the pontine reticular formation ventral to the parabrachial nucleus, not in the parabrachial taste area. Extracellular neural activity was recorded from the parabrachial region in decerebrate rats, and responsiveness to taste stimulation, jaw movements, and electrical stimulation of the superior salivatory nucleus was examined. Neurons responsive to both taste stimulation and antidromic stimulation of the superior salivatory nucleus were found in the pontine reticular formation ventral to the parabrachial nucleus, which responded well to quinine and HCl taste stimuli. Neurons in the parabrachial taste area could respond to four qualities of taste stimulation, but not to antidromic stimulation of the salivary center. These results suggest that aversive taste information from the parabrachial taste area reaches the salivary secretory center via the reticular formation ventral to the parabrachial nucleus.

Animals↗
PubMed1988

Cyclic nucleotides may mediate taste transduction.

Taste stimulus adsorption is believed to occur at the taste cell microvillous membrane. But due to technical difficulties of inserting glass electrodes into the mammalian taste cell, little is known about the mechanisms of taste transduction. Reliable intracellular recordings are necessary to determine the characteristics of taste cells. This has been accomplished previously in the mouse and is reported here. Recent experiments indicated that cyclic nucleotides can act on the inner surface of the membranes of a variety of cells to alter their ion-channel activity, and these substances might act as intracellular transmitters in taste cells. But tight junctions found at the apical membrane of mammalian taste cells do not allow stimuli to enter the taste bud, making it difficult to alter the environment of the taste cell by perfusing with chemical solutions. Here we report that cyclic AMP, cyclic GMP, EGTA or tetraethyl-ammonium electrophoretically injected into the mouse taste cell induce membrane depolarization and increased membrane resistance. These results suggest that a cyclic nucleotide enzymatic cascade, modulated by calcium ions, may mediate the potassium permeability that controls taste, in a way analogous to visual and olfactory transduction.

Animals↗
PubMed2003

The effects of taste and consistency on swallow physiology in younger and older healthy individuals: a surface electromyographic study.

Recent studies done on healthy older individuals have demonstrated elevated taste thresholds for sweet, sour, and salty taste. At suprathreshold levels, such individuals have also shown less ability to discriminate between different intensities of the same tastant. This study was designed to provide information on swallow timing and muscle contraction intensity variables in healthy older people during swallowing of liquid and cottage cheese consistencies in 3 taste conditions (sweet, salty, and sour). Taste and consistency were incorporated in the design of this study to determine not only the effect of taste and consistency independently, but also the effects of taste and consistency in combination. Surface electromyography (EMG) at 3 sites--the orbicularis oris inferior region, submental muscle region, and infrahyoid muscle region--were included in the study. Results revealed that the start of submental muscle activation was significantly later in older participants than in younger participants. The 3 taste conditions had higher EMG levels and shorter activation times than the no-taste condition, although different taste conditions and different muscle sites were affected differently. With thicker consistency, substantially increased EMG amplitude and duration were noted for all three sites. A more pronounced effect of taste was manifested as earlier submental or infrahyoid muscle activation when the three tastants were added to a thicker consistency. Interaction of taste, consistency, and age was also noted for onset time at the submental and infrahyoid sites.

Adolescent↗
PubMed1995

Keratin 18 is associated with a subset of older taste cells in the rat.

All or nearly all intragemmal (elongated) cells of rat taste buds were immunopositive for keratins 7, 8, and 19. In contrast, keratin 18 was detected in 19 +/- 5 cells per taste bud (mean +/- sd), or about 25% of the intragemmal cells. During taste bud development keratins 7, 8, and 19 were evident initially in polygonal cells and later in elongated taste cells. Keratin 8 appeared in vallate taste cells at P0 (postnatal day 0), followed by keratins 7 and 19 at P1, and keratin 18 at P2-P3. Keratin 18 was always limited to elongated cells. The assemblage of elongated taste cells comprising a taste bud began with a single elongated cell, rather than with the synchronous elongation of a cluster of cells. Developmental errors were observed at P2-P3, e.g., some vallate taste cells had a misoriented axis. In order to study the pace of keratin differentiation during cell turnover we injected bromodeoxyuridine (BrdU) into adult rats to monitor taste cell age. Keratin-19-positive intragemmal cells differentiated within 1 day. In contrast, keratin 18 was first detected in cells aged 3 days. Hence, both in taste cell development and replacement, keratin 18 was restricted to the older cells; it was the last taste cell keratin to become expressed during differentiation.

Animals↗
PubMed1998

A novel monoclonal antibody that recognizes apical membrane of frog taste cells.

We established a hybridoma clone 1N1 that produced a monoclonal antibody to stain the apical portion of frog taste cells, by directly immunizing taste discs of the bullfrog (Rana catesbeiana) without any dispersion procedure of the taste organ. The antibody stained discrete regions on the surface of the taste discs, but did not stain the epithelium sheet of the tongue devoid of taste discs. The antibody stained approximately 93% of the taste discs tested (172/184) derived from nine frogs, showing that distribution of the antigen was common to most of the taste discs. The following observations strongly suggested that the antibody recognized a certain antigen on the apical membrane of the taste cells. (i) The antibody selectively stained cross points of intermucus areas on the surface of the taste disc. Neither the mucus cells nor the wing cells that mainly cover the surface were stained with the antibody. (ii) Dispersed taste cells were prepared by calcium ion chelating and subsequently by collagenase treatment to avoid digestion of the antigen. The antibody stained the apical end of the taste cells.

Animals↗
PubMed2000

The effects of sialoadenectomy and exogenous EGF on taste bud morphology and maintenance.

Taste buds on the dorsal tongue surface are continually bathed in saliva rich in epidermal growth factor (EGF). In the following experiment, taste bud number and morphology were monitored following submandibular and sublingual salivary gland removal (sialoadenectomy), to determine if EGF plays a role in the maintenance and formation of taste buds. Adult male rats were divided into four groups: sialoadenectomized (SX, n = 4); sialoadenectomized with EGF replacement (SX + EGF, n = 5); sham-operated (SH, n = 4); and sham-operated with exogenous EGF (SH + EGF, n = 5). After a 3 week recovery, SX + EGF and SH + EGF animals were given 50 microg/day EGF in their drinking water for 14 days. At day 14, saliva was collected, the animals were killed and the presence of EGF determined by radioligand-binding assay. Tongues were removed and histologically examined for the presence and morphology of taste buds on fungiform and circumvallate papillae, or immunostained for the presence of EGF, TGFalpha (transforming growth factor alpha) and EGFR (EGF receptor). The removal of submandibular and sublingual salivary glands resulted in the loss of fungiform taste buds and normal fungiform papillae morphology. These effects were reversed by EGF supplementation, indicating a role for EGF in fungiform taste bud maintenance. In addition, supplementation of EGF to sham-operated animals increased the size of fungiform taste buds. In contrast, removal of salivary glands had no effect on the size, numbers, or morphology of circumvallate taste buds, suggesting that the formation and maintenance of taste buds in fungiform and circumvallate papillae may involve different and distinct processes. EGF, TGFalpha and EGFR were localized to distinct layers of the dorsal epithelium and to within both fungiform and circumvallate taste buds. Their expression within the epithelium or taste buds was not altered with sialoadenectomy, indicating that the actions of endogenous EGF and TGFalpha are distinct and not regulated by exogenous EGF and TGFalpha supplied in saliva.

Animals↗
PubMed2006

Molecular signaling during taste aversion learning.

Behavioral and neural assessment tools have been used to identify cellular and molecular events that occur during taste aversion acquisition. Studies described here include an assessment of taste information processing and taste-illness association using fos-like immunoreactivity (FLI) to mark populations of cells that react strongly to the taste conditioned stimulus (CS), the illness unconditioned stimulus (US), or the pairing of CS and US. Exposure to a novel, but not a familiar, CS taste (saccharin) was found to induce robust increases in FLI in some, but not all, brain regions previously implicated in taste processing or taste aversion learning. Striking effects of taste novelty on FLI were found in central amygdala (CNA) and insular cortex (IC) but not in basolateral amygdala (BLA), pontine parabrachial nucleus (PBN), or nucleus of the solitary tract (NTS). Of those regions responding to taste novelty, only CNA showed significant elevations in FLI in response to the US, LiCl. In additional studies, FLI was examined after an effective training experience, novel CS-US pairing, and compared with an ineffective one, familiar CS-US pairing. After CS-US pairing, taste novelty modulated FLI in virtually all the regions previously implicated in conditioned taste aversion (CTA) learning, including PBN, CNA, BLA, IC, as well as NTS. Thus, a distributed and interdependent neural CTA circuit is mapped using this method, and the use of localized lesion and inactivation studies promises to further define the functional role of structures within this circuit.

Animals↗
PubMed2000

Physiological studies on umami taste.

The first electrophysiological studies on umami taste were conducted with rats and cats. Unlike humans, these animals did not show a large synergism between monosodium glutamate (MSG) and disodium guanylate (GMP) or disodium inosinate (IMP). The taste nerve responses of these animals to umami substances were not differentiated from the salt responses. The canine taste system was sensitive to umami substances and showed a large synergism between MSG and GMP or IMP. The umami substances showed no enhancing effects on other basic tastes. Amiloride, an inhibitor for the response to NaCl, did not inhibit the large response induced by the synergism between MSG and the nucleotides, indicating that the response to the umami substances is independent of the response to salt. Single-fiber analysis on the responses of mouse glossopharyngeal nerve and monkey primary taste cortex neurons also showed that the responses to umami substances are independent of other basic tastes. On the basis of these results, it was proposed that the umami taste is a fifth basic taste, and that there is a unique receptor for umami substances. Hence, we compared the taste of agonists for brain glutamate receptors. In humans, the order of intensity of umami taste induced by a mixture of 0.5 mmol/L GMP and 1.5 mmol/L of various agonists was glutamate > ibotenate > L(+)-2-amino-4-phosphonobutyric acid (L-AP4) = (+/-)1-aminocyclopentane-trans-1,3-dicarboxylic acid (ACPD). Kainate, N-methyl-D-aspartic acid (NMDA) and (RS)-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid (AMPA), which are agonists for ionotropic receptors, had no umami taste. It was concluded that the umami receptor is not identical to any known glutamate receptors; there seems, therefore, to be a unique receptor for umami.

Amino Acids↗
PubMed2000

Receptor and transduction processes for umami taste.

The unique taste of umami argues for a specific receptor at the taste cell level. The taste synergism between monosodium glutamate (MSG) and certain 5'-ribonucleotides provides a pharmacologic test for hypothetical mechanisms of umami taste. Early neurophysiologic and biochemical studies demonstrated specific recognition of L-glutamate by taste tissue and suggested that the synergism found with certain 5'-ribonucleotides was due to a peripheral event. The search for a receptor for umami relies at present on the data in the literature on central nervous system (CNS) glutamate receptors. These data distinguish several classes of receptors on the bases of pharmacologic properties and mode of action. Two hypotheses now seek to explain umami taste transduction. One states that umami is transduced by an N-methyl-D-aspartate (NMDA)-type glutamate ion channel receptor, the other that this taste is transduced via a metabotropic-type glutamate receptor. Evidence for the first hypothesis derives from earlier reconstitution studies, revealing a glutamate-stimulated ion channel conductance whose kinetics were affected by 5'-ribonucleotides. Additional evidence is provided from more recent calcium-imaging and patch-clamp studies, both showing that an ionotropic-type receptor on rodent taste cells mediates glutamate-induced depolarization. Evidence for the second mechanism derives from studies that located the message for an metabotropic-type (mGluR4) receptor to rat taste buds, and from whole-cell patch-clamp recordings that revealed sustained cellular conductances to glutamate and an mGluR4 agonist. It appears likely that both mechanisms are involved in umami taste transduction, suggesting the possibility that reception and transduction of the umami signal constitute a collective property of a number of cells within the taste bud.

Animals↗
PubMed2004

Posttonsillectomy taste distortion: a significant complication.

OBJECTIVE/HYPOTHESIS: Tonsillectomy is among the most commonly performed procedures. As with any surgery, head and neck surgeons must be aware of possible complications and their potential affects. At our smell and taste center, we have been referred several patients in a 6-month period with the complaint of taste distortion after tonsillectomy. We report in this article a patient that complains of taste distortion after a right tonsillectomy for unilateral tonsillar hypertrophy. STUDY DESIGN: A prospective study documenting taste distortion after tonsillectomy using clinical, subjective, and objective evaluation. METHODS: The clinical course of a patient with taste distortion after a tonsillectomy is described. The gustatory function was investigated by conducting electrogustometry and spatial taste testing. Threshold measurements were determined at three left- and three right-side tongue regions: 1) the tongue tip region (innervated by the chorda tympani branch of the facial nerve), 2) the lateral margin of the tongue (anterior to the foliate papillae), and 3) the posterior tongue region (innervated by the lingual branch of the glossopharyngeal nerve). RESULTS: After a complete clinical evaluation and taste testing, it was found that the patient suffered an injury to the right lingual branch of the glossopharyngeal nerve. The close anatomic relationship between the palatine tonsil and lingual branch of the glossopharyngeal nerve makes the nerve vulnerable during tonsillectomy. This injury has caused the patient to suffer ageusia to the right posterior one third of the tongue, compensated by a contralateral phantogeusia (phantom taste) with clinical dysgeusia. The phantogeusia was abolished by application of anesthetic to the area where the phantom was perceived. We propose that the phantogeusia is the result of release-of-inhibition in the contralateral glossopharyngeal nerve. CONCLUSION: Taste distortion (including, phantogeusia and dysgeusia) after tonsillectomy is rarely reported as a complication but has a significant impact on quality of life. This article examines the taste distortion presence as a complication after tonsillectomy to make head and neck surgeons aware of this serious complication and the pathophysiology of taste distortion.

Dysgeusia↗
PubMed1996

Clinical and physiological investigations in patients with taste abnormality.

The cause of taste abnormality was investigated in 25 patients with decreased taste sensation (hypogeusia group) and 14 patients with abnormal taste sensation (dysgeusia group) by examining taste threshold, salivary flow rate, Candida cell culture, and laboratory examination of peripheral blood. The cause of hypogeusia was identified as iron deficiency in 7 patients, oral candidiasis in 6, hyposalivation (xerostomia) in 6, and psychiatric distress in 3, and could not be determined in 3 (idiopathic). Dysgeusia was associated with psychiatric distress in 8 patients, oral candidiasis in 3, drug medication in 2, and hyposalivation in 1. In the hypogeusia group, the decreased taste sensation generally corresponded with elevated taste thresholds, which decreased along with improvement of the decreased taste sensation in all except the 3 patients with psychiatric etiology and 2 of the 3 patients with idiopathic etiology. In contrast, no elevation or depression of taste thresholds were observed in the dysgeusia group, and the abnormal taste sensation did not disappear in most cases; however, drug-induced dysgeusia improved completely within 2 months after cessation of the drug administration. The serum copper and zinc levels were not decreased in any patient, but a decreased serum iron level was observed in 7 patients. Based on these results, it is concluded that abnormal taste sensation may be induced by many oral and systemic disturbances and that hypogeusia, which may be induced by deficiency of iron but not of zinc or copper, is usually accompanied by elevation of taste thresholds, while dysgeusia is not.

Adult↗
PubMed1998

Quantitative relationships between taste bud development and gustatory ganglion cells.

To determine whether patterns of taste bud innervation change during postnatal rat development, the number of geniculate ganglion cells that innervate single taste buds were quantified in adult and developing rats. While there was a large variation in numbers of ganglion cells that innervate individual taste buds, there was a high degree of organization in the system. Namely, the number of labeled geniculate ganglion cells innervating a taste bud was highly correlated with the size of the taste bud. This relationship between taste bud size and number of innervating ganglion cells develops over a prolonged postnatal period and is not established until postnatal day 40 (P40), when taste buds reach their adult size. In a second series of experiments, we sought to determine whether neural rearrangement of chorda tympani neurons is responsible for the development of this relationship by double-labeling single taste buds at different ages. We found that the number of ganglion cells innervating individual taste buds on P10 predicts the size that taste buds become by P40. This finding suggests that neural rearrangement is not responsible for establishing the relationship between taste bud size and the number of innervating ganglion cells during development. More importantly, it strongly suggests that the 'neural template' for the mature innervation pattern is determined during early postnatal development.

Animals↗