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PubMed1988

Critical periods in the development of motoneurons.

We define "critical periods" in motoneuron development as periods when neurons can express a particular aspect of their sequence of differentiation only within a limited time interval. Embryonic rat motoneurons were axotomized and their survival and capacity to regenerate their axons studied at later times. A critical period of absolute target dependence extended from embryonic day 14 (E14) until E17. All motoneurons axotomized during this interval died, while cells axotomized earlier or later could survive. During the period E18- postnatal day 14 (PN14) regenerating motoneurons accurately reinnervated motor unit territories of normal size, and intact motoneurons did not exhibit collateral sprouting in response to denervation of adjacent muscle fibres. After PN21, muscle fibres were reinnervated in a spatially indiscriminate manner, and collateral sprouting occurred in response to partial denervation of muscles. We conclude that adult motoneurons have a powerful capacity to reinnervate denervated muscles, even in an inappropriate manner, in contrast to motoneurons in neonates which can recapitulate embryogenesis and accurately reinnervate their proper motor unit territories.

Animals↗
PubMed2007

Adolescent cortical development: a critical period of vulnerability for addiction.

Cortical growth and remodeling continues from birth through youth and adolescence to stable adult levels changing slowly into senescence. There are critical periods of cortical development when specific experiences drive major synaptic rearrangements and learning that only occur during the critical period. For example, visual cortex is characterized by a critical period of plasticity involved in establishing visual acuity. Adolescence is defined by characteristic behaviors that include high levels of risk taking, exploration, novelty and sensation seeking, social interaction and play behaviors. In addition, adolescence is the final period of development of the adult during which talents, reasoning and complex adult behaviors mature. This maturation of behaviors corresponds with periods of marked changes in neurogenesis, cortical synaptic remodeling, neurotransmitter receptors and transporters, as well as major changes in hormones. Frontal cortical development is later in adolescence and likely contributes to refinement of reasoning, goal and priority setting, impulse control and evaluating long and short term rewards. Adolescent humans have high levels of binge drinking and experimentation with other drugs. This review presents findings supporting adolescence as a critical period of cortical development important for establishing life long adult characteristics that are disrupted by alcohol and drug use.

Adolescent↗
PubMed2005

Rapid eye movement sleep deprivation in post-critical period, adolescent rats alters the balance between inhibitory and excitatory mechanisms in visual cortex.

Suppression of rapid eye movement sleep (REMS) in developing animals has both anatomical and physiological consequences. We have recently shown that initiating REMS deprivation (REMSD) prior to the end of the critical period in young rats delays termination of the critical period (CP) in visual cortex, and, consequently, the synaptic plasticity mechanisms that support a developmentally regulated form of long-term potentiation (LTP) are maintained in an immature state [J.P. Shaffery, C.M. Sinton, G. Bisset, H.P. Roffwarg, G.A. Marks, Rapid eye movement sleep deprivation modifies expression of long-term potentiation in visual cortex of immature rats, Neuroscience, 110 (2002) 431-443]. In CP animals, high-frequency, theta burst stimulation (TBS) directed at the white matter (WM) below visual cortex produces LTP in the post-synaptic cells in layer II/III (LTPWM-III). However, LTPWM-III can be induced in cortical tissue taken from REMS-deprived animals for up to a week beyond the usual end of the CP [J.P. Shaffery, C.M. Sinton, G. Bisset, H.P. Roffwarg, G.A. Marks, Rapid eye movement sleep deprivation modifies expression of long-term potentiation in visual cortex of immature rats, Neuroscience, 110 (2002) 431-443]. Further, in post-CP, adolescent animals (as late as postnatal day 60), REMSD appears to unmask synaptic plasticity mechanisms that allow for production of developmentally regulated LTPWM-III [J.P. Shaffery, J. Lopez, G. Bissette, H.P. Roffwarg, Rapid eye movement sleep deprivation revives a form of developmentally regulated synaptic plasticity in the visual cortex of post-critical period rats, Neurosci Lett., (2005), in press]. It has been proposed that REMSD's effects on production of LTPWM-III result from a reduction in efficiency of the inhibitory mechanisms thought to precipitate termination of the CP of brain development [J.P. Shaffery, J. Lopez, G. Bissette, H.P. Roffwarg, Rapid eye movement sleep deprivation revives a form of developmentally regulated synaptic plasticity in the visual cortex of post-critical period rats, Neurosci Lett., (2005), in press]. In this study we tested the hypothesis that low-frequency stimulation (LFS) of the fibers of the WM, which usually produces the related form of synaptic plasticity, long-term depression (LTD), will also reflect the reduction in inhibitory tone. We report here that LFS protocols, which in normally sleeping, adolescent rats usually produce either LTD or no change in response magnitude, in REMS-deprived, adolescent rats are more likely to produce LTP.

Age Factors↗
PubMed2003

Decline of the critical period of visual plasticity is concurrent with the reduction of NR2B subunit of the synaptic NMDA receptor in layer 4.

The specific composition of NMDA receptor subunits is thought to underlie the developmental plasticity of the cortex revealed by unbalanced binocular stimulation. However, evidence that NR2 subunits change in correlation with the critical period at locations that are relevant to visual plasticity has been missing. Using preembedding and postembedding immunostaining, as well as electron microscopy, we quantified the volumetric densities of NR1-, NR2A-, and NR2B-containing synapses in layers 4 and 2/3 of the ferret visual cortex at different postnatal ages. Before eye opening, NR2A is encountered infrequently at postsynaptic sites in layer 4, but it increases sharply by postnatal day 34. In the subsequent weeks, postsynaptic NR2A labeling increases gradually in both layers 4 and 2/3 to become the most prevalent subunit in the adult animal. The NR2B subunit is the more prevalent subunit at the onset of the critical period of cortical plasticity. However, it displays different developmental patterns in layers 4 and 2/3. Although no change occurs in synaptic NR2B density in layer 2/3, in layer 4, NR2B maintains its high levels through the peak of the critical period and then becomes significantly reduced by the end of the peak of the critical period. This low level is maintained throughout adulthood. Our results demonstrate a correlation between the loss of NR2B subunits from layer 4 synaptic sites and the decline of the critical period, suggesting that the presence of NR2B subunits at synaptic sites could be a permissive factor regulating the ocular dominance plasticity of the developing cortex.

Age Factors↗
PubMed2000

Neurotrophin-4/5 alters responses and blocks the effect of monocular deprivation in cat visual cortex during the critical period.

The mechanisms underlying changes in neural responses and connections in the visual cortex may be studied by occluding one eye during a critical period in early postnatal life. Under these conditions, neurons in the visual cortex rapidly lose their responses to the deprived eye and ultimately lose many of their inputs from that eye. Cats at the peak of the critical period received infusions of exogenous neurotrophin NT-4/5 into primary visual cortex beginning before a short period of monocular deprivation. Within areas affected by NT-4/5, cortical cells remained responsive to the deprived eye, and maps of ocular dominance were no longer evident using intrinsic-signal optical imaging. Cortical cells also became broadly tuned for stimulus orientation and less responsive to visual stimulation through either eye. These effects required at least 48 hr exposure to the neurotrophin and were specific for trkB, because they were not seen with the trkA or trkC ligands NGF or NT-3. Even after neurons had already lost their responses to the deprived eye, subsequent NT-4/5 infusion could restore them. The NT-4/5 effects were not seen after the critical period. Together, these results suggest that trkB activation during the critical period may promote promiscuous connections independent of correlated activity.

Aging↗
PubMed2003

The freshman year of college as a critical period for weight gain: an initial evaluation.

Prevention of obesity and weight maintenance have become important public health issues. One strategy for prevention of obesity is to identify critical periods of weight gain across the life span. The purpose of this initial evaluation was to determine whether the transition from high school to college is such a critical period. A total of 135 college students were weighed in September and December of their freshman year and a subset also provided data in May. Results showed that statistically significant but modest weight increases occurred during the freshman year for most participants. However, one quarter of participants gained at least 2.3 kg during the first semester of college, and the proportion of participants classified as overweight or obese increased markedly. For this subset of participants, the freshman year of college could be considered a critical period for weight gain. Identifying critical periods for weight gain may be an important first step towards the development of effective obesity prevention programs.

Journal Article↗
PubMed2000

Critical periods during sensory development.

Recent studies have made progress in characterizing the determinants of critical periods for experience-dependent plasticity. They highlight the role of neurotrophins, NMDA receptors and GABAergic inhibition. In particular, genetic manipulation of a single molecule, brain-derived neurotrophic factor (BDNF), has been shown to alter the timing of the critical period of plasticity in mouse visual cortex, establishing a causal relation between neurotrophin action, the development of visual function, and the duration of the critical period.

Animals↗
PubMed1984

Critical periods for effects of monocular deprivation: differences between striate and extrastriate cortex.

The critical period of susceptibility to effects of monocular deprivation was compared in striate cortex and the lateral suprasylvian (LS) visual area of cortex. Twenty-three cats received monocular lid suture for a period of 4 weeks beginning at 4, 12, 18, 26, or 35 weeks of age or as adults. Immediately following the deprivation, single cell recordings were carried out in both cortical areas of each cat. Recordings also were made from five normally reared control cats. For both striate and LS cortex, early monocular deprivation had marked effects on neuronal ocular dominance, including an increased percentage of cells dominated by the nondeprived eye, a decreased percentage of cells dominated by the deprived eye, and a decreased percentage of binocularly driven cells. In both cortical areas, these effects were maximal in animals deprived at 4 weeks of age. Both areas then showed similar monotonic declines in effects of the deprivation following onsets from 4 to 18 weeks of age. However, in older animals there were clear differences in the effects of monocular deprivation on LS and striate cortex. In LS cortex, the monotonic decline in effects continued until 26 weeks of age, and no significant abnormalities were present in animals deprived at 26 weeks of age or older. In striate cortex, however, the effects of monocular deprivation remained relatively constant following onsets from 18 to 35 weeks of age, and significant abnormalities in all measures of ocular dominance were present when deprivation was begun as late as 35 weeks of age. Within-animal comparisons indicated that the greater effects of monocular deprivation on striate cortex than on LS cortex were present in every cat deprived at 26 or 35 weeks of age. Neither cortical area showed significant abnormalities following monocular deprivation in adult cats. These results indicate that the critical period for effects of the same regime of monocular deprivation is over sooner in LS cortex (between 18 and 26 weeks of age) than in striate cortex (after 35 weeks of age). This observation has important implications for an understanding of the sites and mechanisms of effects of visual deprivation and the mechanisms that control critical periods of development.

Animals↗
PubMed1985

The critical period for corpus callosum section to affect cortical binocularity.

The period of time during which surgical section of the corpus callosum (CC) is effective in altering the physiological properties of cells in cat striate cortex was investigated. Cats which had the CC transected between 13 days and 24 weeks of age were studied using extracellular, single-unit recording procedures. Analysis of the results from 1,747 cortical units indicate that when the CC was sectioned prior to 19 days of age there was a reduction in the encounter rate of binocularly activated neurons and an increase in the proportion of neurons dominated by the contralateral eye. The decrease in cortical binocularity was observed in both simple and complex cell populations, and at all receptive field eccentricities studied (0-39 degrees). However, when the CC was sectioned after 19 postnatal days, no physiological changes were detected. Thus, in contrast with previous studies (Payne et al. 1980a, b) no changes were found following CC section in adult cats. The results therefore define a critical period which ends before 3 weeks of age during which corpus callosum section reduces striate cortex binocularity. Although the corpus callosum critical period is much shorter than the critical period for experiential alterations in cortical binocularity, the physiologically determined limits of the callosal critical period agree with the behaviorally determined limits previously found for the callosal critical period (Elberger 1984).

Animals↗
PubMed1985

Postnatal developmental profiles of filamentous actin and of 200 kDa neurofilament polypeptide in the visual cortex of light- and dark-reared rats and their relationship to critical period plasticity.

The concept of critical period plasticity in rat visual cortex has been studied in terms of changes in the level of filamentous actin and of the 200 kDa neurofilament polypeptide. Our results suggest that the postnatal developmental profile of filamentous actin is affected by visual experience, as a consequence of eye-opening. No such correlation, however, is detected for the 200 kDa neurofilament polypeptide. The significance of these findings in relationship to neuronal plasticity is discussed in terms of changes in the state and equilibrium conditions of the cytoskeletal proteins.

Actins↗
PubMed2003

Laminar distribution of NMDA receptor subunit (NR1, NR2A, NR2B) expression during the critical period in cat visual cortex.

Changes in NMDA subunit composition may be part of the molecular basis for critical period plasticity. The present study used immunohistochemistry to determine developmental changes in the laminar distribution of the three major cortical NMDA subunits (NR2A, NR2B, NR1) during the critical period in cat visual cortex. For all three subunits, at 1 week staining was concentrated in two bands: an upper band consisting of layer I, the compact zone and the upper half of the cortical plate; a lower band consisting of layers V and VI. In the lower part of the cortical plate (immature layer IV) staining was very low. For NR2A and NR2B, immunoreactivity in layer IV remained low until 10 weeks of age. At 20 weeks and adult, layer IV filled in and NR2A and NR2B label was rather uniform across all layers. NR1 showed a developmental pattern of expression different from NR2A and NR2B after 1 week. At 5 and 10 weeks, label was prominent in layer IV and superficial layers, but low in layers V and VI. The main change after 10 weeks was a progressive decrease in staining, such that in older animals label was markedly densest in superficial layers. Thus, during the rise of the critical period, NR1 is the dominant subtype in layer IV and could play a role in anatomical ocular dominance column formation and plasticity. At the same time, NR2A and NR2B subunits are concentrated outside layer IV, and could be related to physiological plasticity in extragranular layers, which precedes and outlasts plasticity in layer IV. For all three NMDA receptor subunits, the laminar distribution was similar in normal and dark reared visual cortex at 20 weeks, indicating that the developmental changes in laminar pattern of expression are independent of visual input.

Afferent Pathways↗
PubMed1995

A revised critical period for the sexual differentiation of the sexually dimorphic nucleus of the preoptic area in the rat.

The volume of the sexually dimorphic nucleus of the preoptic area (SDN-POA) of the rat is several times larger in males than in females. Several studies have established the importance of gonadal steroids perinatally in the sexual differentiation of the SDN-POA as well as a critical period for the permanent influences of exogenous androgen on the volume of the nucleus. Recent preliminary evidence from this laboratory had suggested, however, that the critical period for the effects of the removal of endogenous gonadal steroids on SDN-POA volume may not match that for the administration of exogenous gonadal steroids. A series of experiments was designed to examine further the effects of the removal of endogenous gonadal steroids on adult SDN-POA volume by castrating male rats at various ages. In spite of a rather clear definition of a postnatal critical period for the effects of exogenous steroid administration, the results of this study indicate that the volume of the SDN-POA is sensitive to the removal of endogenous gonadal steroids for a prolonged period of time, extending through at least day 29 postnatally. The data suggest that there may be multiple critical periods for the sexual differentiation of the SDN-POA and reinforce the concept that these critical periods are distinct from those for other sexually differentiated parameters.

Animals↗
PubMed1997

Synelfin regulation during the critical period for song learning in normal and isolated juvenile zebra finches.

Male zebra finches (Taeniopygia guttata) learn to sing during a critical period in adolescence. We previously described a presynaptic protein, synelfin, whose mRNA is increased early in this critical period in a brain nucleus specifically implicated in song learning, lateral MAN (lMAN). In the current study, in situ hybridization was used to map this change in gene expression to the subregion of lMAN that projects to the robust nucleus of the archistriatum (RA), the principal motor output of the telencephalic circuit that controls song production. Using confocal immunofluorescence microscopy, we detected numerous puncta of synelfin immunoreactivity that apparently represent presynaptic terminals from lMAN in the RA of young males. Synelfin immunoreactivity in RA declined abruptly between 40 and 45 days of age, a time of major synaptic reorganization in RA. This change did not occur until about 10 days after the decline in synelfin mRNA in cell bodies within lMAN, indicating a relatively slow turnover of the protein in presynaptic terminals and suggesting that some of the functional changes that occur during the critical period may arise from regulatory decisions that were initiated a week or more earlier. Depriving birds of tutoring did not halt or delay the decline of synelfin mRNA in lMAN. This change in gene expression must not be a consequence of early song learning, but may reflect an innate or programmed step in song circuit development.

Amyloid↗
PubMed2001

Critical periods for the effects of alcohol exposure on learning in rats.

Critical periods for alcohol-induced deficits in spatial navigation and passive avoidance learning were investigated with a rat model of fetal alcohol syndrome. Rats were exposed to alcohol prenatally (Gestational Days 1-10 or 11-22) or postnatally (Postnatal Days 2-10) or throughout all 3 periods. Offspring were tested in either a spatial navigation or an avoidance task as juveniles or adults. As juveniles, the combined exposure group took longer to learn the spatial navigation task compared with all other groups. This effect was not seen in adults. Passive avoidance performance was not affected. These results suggest that long-term exposure to alcohol during development has adverse effects on spatial learning. The lack of differences in the short-term exposure groups implies that there may not be 1 critical period of alcohol exposure, but that the adverse effects of alcohol during development may be cumulative on some behaviors.

Age Factors↗
PubMed2004

"Critical periods" in vestibular development or adaptation of gravity sensory systems to altered gravitational conditions?

1. A feature of sensory, neuronal and motor systems is the existence of a critical period during their development. Modification of environmental conditions during this specific period of life affects development in a long-term manner, or even irreversibly. Deprivation is the prefered approach to study the existence and duration of critical periods. For gravity sensory systems, space flights offer the only opportunity for deprivation conditions. 2. Studies in a fish (Oreochromis mossambicus) and an amphibian (Xenopus laevis) revealed a significant sensitivity of their roll-induced static vestibuloocular reflex (rVOR) to a 9- to 10-day gravity deprivation (microgravity) during a spaceflight. In some instances, the rVOR was augmented after the flight as demonstrated in young Oreochromis which were launched when their rVOR had not been developed, and in Xenopus tadpoles launched after their rVOR had developed. Fish which could perform the rVOR at launch were insensitive to microgravity exposure. A similar insensitivity to microgravity was observed in Xenopus tadpoles with normal body shape which had not yet developed their rVOR at launch. Some tadpoles, however, developed an upward bended tail during their space flight; their rVOR was significantly depressed after termination of microgravity independent of the age at onset of the flight. Hypergravity depressed the rVOR for all so far tested developmental stages in both Oreochromis and Xenopus. 3. Both adaptive processes during exposure to altered gravity as well as the existence of a critical period in vestibular development might be responsible for the modulation of the rVOR recorded after exposure to altered gravity. Deprivation studies have to be extended to older developmental stages to test the possibility of a critical period; however, this approach is limited due to the low number of space flights.

Adaptation, Physiological↗
PubMed2003

Critical evidence: a test of the critical-period hypothesis for second-language acquisition.

The critical-period hypothesis for second-language acquisition was tested on data from the 1990 U.S. Census using responses from 2.3 million immigrants with Spanish or Chinese language backgrounds. The analyses tested a key prediction of the hypothesis, namely, that the line regressing second-language attainment on age of immigration would be markedly different on either side of the critical-age point. Predictions tested were that there would be a difference in slope, a difference in the mean while controlling for slope, or both. The results showed large linear effects for level of education and for age of immigration, but a negligible amount of additional variance was accounted for when the parameters for difference in slope and difference in means were estimated. Thus, the pattern of decline in second-language acquisition failed to produce the discontinuity that is an essential hallmark of a critical period.

Age Factors↗
PubMed1981

In vitro resetting of the circadian clock in the Aplysia eye. II. The critical period for optic nerve activity.

When constant light is 18, 21, or 24 hr, a constant light to constant darkness (LL/DD) transition in vitro results in a phase difference (delta psi) between an eye that is attached to the cerebral ganglion and its partner that is detached by cutting its optic nerve just prior to the LL/DD. This delta psi develops during the first 24 hr after LL/DD as the result of efferent activity in the optic nerve of the attached eye. By cutting the nerve of the original attached eye at various times after LL/DD, we determined when the phase resetting nerve activity occurs. This experiment was carried out following LL = 18, 21, and 24 hr. Optic nerve activity at the time of and for several hours after LL/DD did not produce nerve-dependent resetting (NDR). Instead, there was a restricted critical period, 2 to 3 hr long during which the effective nerve activity occurred. Following 24 hr of LL, a 2-hr window of optic nerve activity during the critical period could produce NDR, but a 2-hr window outside of the critical period was ineffective. The resetting effect of nerve activity was produced suddenly as if NDR were an all-or-nothing event. Following 18, 21, and 24 hr of LL, the critical period occurred 10, 7, and 4 hr, respectively, after LL/DD. In each case, this was about 28 hr after the last dawn of the light cycles to which the animal had been exposed prior to LL. We conclude that the critical period is timed by a mechanism that is not reset by LL/DD.

Action Potentials↗
PubMed2004

Different dopamine D1 and D2 receptors expression after motor activity in the striatal critical period.

Circling training during rat striatum postnatal critical period (PN30 to 37 days) induces a life-lasting co-reduction of muscarinic acetylcholine receptors (mAChR) and dopamine D2 receptors (D2R) binding. Here, we evaluated the expression of D1R and D2R under similar experimental conditions. Trained rats showed a decrease of 40% in D2R binding sites (p<0.01) and of 45% in the D2R mRNA expression which involve short (p<0.05) and long (p<0.01) isoforms. In contrast, D1R binding sites nor its mRNA expression levels were affected by training, indicating a differential synaptic refinement during this ontogenetically fixed period.

Animals↗