The effects of antioxidants in the senescent auditory cortex.

We investigated whether a 2-month dietary supplementation of antioxidants, in the form of blueberry phytochemicals, could reverse or retard the age-related decline in temporal processing speed observed in the aged rat. To this end, extracellular single unit responses to frequency modulated (FM) sweeps were recorded in the primary auditory cortex (AI) of aged rats that had been placed on either a blueberry-supplemented or control diet 2 months prior to the physiological recordings. Results showed that most cells recorded from the blueberry-fed rats responded most vigorously to fast FM sweeps, similar to that observed in young rats. In contrast, the majority of cells recorded from the control rats showed a preference for slow FM sweep rates. These results suggest that age-related changes in temporal processing speed in A1 may be reversed by dietary supplementation of blueberry phytochemicals.

The effects of aging in the medial geniculate nucleus: a comparison with the inferior colliculus and auditory cortex.

A common problem among the elderly is a difficulty in discriminating speech. One factor that may contribute to this is deterioration in the ability to process the dynamic components of speech such as formant transitions. The frequency-modulated (FM) sweep is a useful stimulus for investigating the neural basis of temporal processing speed since it has features in common with formant transitions. Previously, we showed that when cells in the auditory cortex of aged animals were presented with FM sweeps, they exhibited a decrease in temporal processing speed when compared to cells recorded from young animals. However, this was not the case for cells in the inferior colliculus (IC) where neural responses did not appear to be affected by aging. One question that remains is how the auditory thalamus is affected by aging: Is it similar to that of the auditory cortex or of the IC. To this end, single units were recorded from the ventral division of the medial geniculate nucleus (MGNv) of young and aged anaesthetized rats in response to FM sweeps. Results showed that there were no age-related differences in speed or direction selectivity of FM sweep responses in the MGNv. When compared with units recorded from the IC and AI, the responses of MGNv neurons were similar to those of the IC. This suggests that temporal processing speed is affected by aging in the cortex, but not in the auditory thalamus or midbrain.

Factors associated with the catastrophic decline of a cloudforest frog fauna in Guatemala.

Comparison of recent and historical surveys of frog populations in cloudforest habitat in Sierra de las Minas, Guatemala, indicated population declines and local extirpation of several species. Pathological exams of diseased tadpoles indicated infection by amphibian chytridiomycosis. The local habitat has been severely altered by recent establishment of large-scale leatherleaf fern production. Analysis of water chemistry at our study site suggested increased nitrogenation associated with the leatherleaf industry.

Frequency modulated sweep responses in the medial geniculate nucleus.

A basic feature of communication signals is a dynamic change in frequency. One stimulus that lends itself well to investigating the frequency changes contained in these signals is the frequency modulated (FM) sweep. While many studies have investigated FM sweep responses in the auditory midbrain and cortex, relatively few have examined them in the thalamus. To this end, we investigated the responses of single units in the ventral division of the medial geniculate nucleus (MGNv) of the rat to FM sweeps. Both upward- (changing from low to high frequency) and downward-directed (changing from high to low frequency) FM sweeps were presented at four rates of frequency modulation (i.e., speed). Results showed that the majority (76%) of the cells preferred fast or medium FM sweeps. For direction selectivity, just under half of the units (47%) exhibited a preference for the direction of FM sweep. The results suggest that there is a greater degree of direction but not speed selectivity at progressively higher levels in the auditory pathway.

Temporal processing speed in the inferior colliculus of young and aged rats.

A common problem among the elderly is a difficulty in discriminating speech. One factor that may contribute to this is a deterioration in the ability to process dynamic aspects of speech such as formant transitions. Recently, Mendelson and Ricketts [Mendelson, J.R., Ricketts, C., Hear. Res. 158 (2001) 84-94] showed that cells recorded from the auditory cortex of aged animals exhibited a decrease in temporal processing speed compared to young animals. In the present study, we examined whether this age-related effect was exclusive to the auditory cortex or whether it was apparent subcortically. To this end, single units were recorded from the inferior colliculus (IC) of young and aged rats in response to frequency modulated (FM) sweeps. Results showed that there was no age-related difference in speed or direction selectivity of FM sweep responses in the IC. The present results suggest that the effect of aging on temporal processing speed occurs in the cortex, but not subcortically.

Age-related changes in the visual cortex.

The ability to accurately perceive the speed of moving objects is one of many visual functions that decline with age. One factor that may contribute to this is a deterioration in temporal processing speed. At present, there is a dearth of information concerning how this may occur in the central nervous system, particularly in the visual cortex. Thus, in the present study, we investigated the neural basis of speed and temporal processing in areas 17 and 18 of visual cortex in young and aged rats using either a moving bar of light or a series of flashing lights. Our results showed that the mean preferred speed of a moving bar of light was significantly reduced in aged as compared to young animals. We also found that cells recorded from young animals were able to entrain to a higher frequency of flashing light stimuli than those recorded from aged animals. In addition, we found no age-related differences between cortical fields. These results suggest an age-related difference in temporal processing speed at the level of visual cortex.

Critical flicker frequency responses in visual cortex.

Critical flicker frequency (CFF) threshold is defined as the frequency at which a flickering light is indistinguishable from a steady, non-flickering light. CFF is useful for assessing the temporal characteristics of the visual system. While CFF responses are believed to reflect activity in the central visual system, little is known about how these temporal frequencies are processed in the visual cortex. The current paper estimated the CFF threshold for cells in the rat visual cortex by recording single unit responses to flickering stimuli. Results showed that: (1) there was a broad range of temporal tuning, (2) CFF threshold was lower in simple cells than in complex and hypercomplex cells, and (3) there was no significant difference in CFF threshold between areas 17 and 18.

Age-related temporal processing speed deterioration in auditory cortex.

A common problem among the elderly is a difficulty in discriminating speech sounds. One factor that may contribute to this is a deterioration in the ability to process dynamic aspects of speech such as formant transitions. For the aging auditory system, this deterioration in temporal processing speed may be manifest as a deficit in encoding time-varying sounds that contain rapidly changing frequencies such as formant transitions. The primary goal of this study was to explore the neural basis of the effects of aging on temporal processing speed. To this end, single units were recorded from the auditory cortex of young and aged rats in response to frequency-modulated (FM) sweeps that changed from trial to trial in both direction and speed. Results showed that the majority of cells recorded from young rats responded most vigorously to fast and medium speeds. By contrast, the majority of units recorded from aged animals responded best to slow speeds. For preferred direction of FM sweep, similar results were observed for both age groups, namely, approximately half of the units exhibited a direction-selective response. The results of the present study demonstrate an age-related decrease in the rate of change of frequency that can be processed by the auditory cortex.

Phylogeography and speciation of the morphologically variable, widespread species Bufo valliceps, based on molecular evidence from mtDNA.

The widespread, lowland toad Bufo valliceps has an unusual distribution in North and Middle America that straddles two major biogeographic areas; previous morphological studies of this species suggested the existence of two species. We used mitochondrial DNA sequences to examine the phylogeography of this species and discovered the existence of two distinct clades. We recognize these as two species: B. valliceps and B. nebulifer. These molecular data support morphological data from previous studies. Our results show low levels of molecular variation in a morphologically uniform temperate species (B. nebulifer) and high levels of molecular variation in a morphologically variable tropical species (B. valliceps), providing an example of molecules matching morphology. Two biogeographic hypotheses are tested to explain the current distribution of these species, based on a calibrated rate of evolution and the percent sequence divergence between the two species. A more recent Pleistocene dispersal event, followed by vicariance associated with rising sea level, is rejected in favor of an earlier Miocene-Pliocene vicariant hypothesis associated with the formation of the Trans-Mexican Neovolcanic Belt.

Auditory cortical responses to the interactive effects of interaural intensity disparities and frequency.

Under natural conditions, stimuli reaching the two ears contain multiple acoustic components. Rarely does a stimulus containing only one component (e.g. pure tone burst) exist outside the realm of the laboratory. For example, in sound localization the simultaneous presence of multiple cues (spectral content, level, phase, etc.) serves to increase the number of available cues and provide the listener with more information, thereby helping to reduce errors in locating the sound source. The present study was designed to explore the relationship between two acoustic parameters: stimulus frequency and interaural intensity disparities (IIDs). By varying both stimulus frequency and IIDs for each cell, we hoped to gain insight into how multiple cues are processed. To this end, we examined the responses of neurons in cat primary auditory cortex (AI) to determine if their sensitivity to IIDs changed as a function of stimulus frequency. IIDs ranging from +30 to -30 dB were presented at different frequencies (frequency was always the same in the two ears). We found that approximately half of the units examined exhibited responses to IIDs that varied as a function of stimulus frequency (i.e. displayed some form of IID x Freq dependency). The remaining units displayed IID responses that were not clearly related to stimulus frequency.

Responses to time-varying stimuli in rat auditory cortex.

Responses to frequency modulated (FM) sweeps were recorded in rat primary auditory cortex. Forty-four percent of the cells were direction-selective. For speed selectivity, the majority of the cells preferred faster sweeps. The results suggest that rat auditory cortex may be used for processing communication signals of their predators or for detecting spectral changes in acoustic signals.

Functional topography of cat primary auditory cortex: response latencies.

Minimum onset latency (Lmin) of single- and multiple-unit responses were mapped in the primary auditory cortex (AI) of barbiturate-anesthetized cats. Contralateral Lmin for multiple units was non-homogeneously distributed along the dorso-ventral/isofrequency axis of the AI. Responses with shorter latencies were more often located in the central, more sharply tuned region while longer latencies were more frequently encountered in the dorsal and ventral portions of the AI. For single units, a large scatter of Lmin values was found throughout the extent of the AI including cortical depth. The relationship between Lmin and previously reported spectral, intensity and temporal parameters was analyzed and revealed statistically significant correlations between minimum onset latency and the following response properties in some but not all studied animals: sharpness of tuning of a frequency response area 10 dB above threshold, broadband transient response, strongest response level, monotonicity of rate/level functions, dynamic range, and preferred frequency modulation sweep direction. This analysis suggests that Lmin is determined by several independent factors and that the prediction of Lmin based on relationships with other spectral and temporal response properties is inherently weak. The spatial distribution and the functional relationship between these response parameters may provide an important aspect of the time-based cortical representation of specific features in the animal's natural environment.

Alterations in visual receptive fields in the superior colliculus induced by amphetamine.

Visual response properties were examined in the superficial layers of the superior colliculus (SC) of anesthetized, paralyzed cats before and after i.v. administration of d-amphetamine. Receptive fields (RFs) of single SC units were plotted using small spots of light presented to the contralateral eye. Within the first hour following d-amphetamine injections, RF size gradually increased, reaching a maximum 86 min post-injection. On average, the area of the RF increased by 5.6 times and RF expansion was observed in all single units examined in the superficial layers. Over the subsequent 4-8 h following the injection, RF area gradually decreased and returned to control dimensions. Most RFs displayed asymmetrical patterns of expansion, showing relatively more horizontal than vertical growth. As RF expansion developed, responses to stimuli flashed "on" and "off" at various locations both inside and outside the borders of the control RF became progressively more vigorous. In contrast, no significant changes were noted in direction-selective responses at any time after d-amphetamine injections. Using an array of light bar stimuli of different lengths, the strength of surround suppression was found to be significantly diminished by d-amphetamine. The reduction in surround suppression was especially clear for bar lengths which exceeded the diameter of the control RF. No RF expansion was observed in the superficial layers of the SC when d-amphetamine was injected intravitreally. Furthermore, d-amphetamine had no discernable effect on the RF sizes of cells in the visual cortex. These results suggest that the RF changes in the SC were not of either retinal or cortical origin. We conclude that the mean retinal area which can potentially influence the activity of RFs in the superficial layers of the SC may be on average over 5 times greater than the RF area determined using conventional methods and criteria. These findings raise the interesting possibility that the relatively small size and sharp borders characteristic of RFs in the superficial layers arise from local inhibitory networks which delimit a broader field of excitatory activity supplied by retinal and cortical afferent terminals. Thus, in order to generate the RF changes observed here, either these local inhibitory circuits are amphetamine sensitive, or more likely, these inhibitory networks are dynamically modulated by an, as yet unidentified, amphetamine-sensitive input affecting visual RFs in the superficial layers.

Functional topography of cat primary auditory cortex: responses to frequency-modulated sweeps.

The spatial distribution of neuronal responses to frequency-modulated (FM) sweeps was mapped with microelectrodes in the primary auditory cortex (AI) of barbiturate-anesthetized cats. Increasing and decreasing FM sweeps (upward- and downward-directed FM sweeps, respectively) covering a range of 0.25-64.0 kHz were presented at three different rates of frequency change over time (i.e, sweep speed). Using multiunit recordings, the high-frequency domain (between 3.2 and 26.3 kHz) of AI was mapped over most of its dorsoventral extent (as determined by the distribution of the excitatory bandwidth, Q10dB) for all six cases studied. The spatial distributions of the preferred sweep speed and the preferred sweep direction were determined for each case. Neuronal responses for frequency sweeps of different speeds appeared to be systematically distributed along the dorsoventral axis of AI. In the dorsal region, cortical cells typically responded best to fast and/or medium FM sweeps, followed more ventrally by cells that responded best to medium--then slow--, then medium-speed FM sweeps. In the more ventral aspect of AI (which in some cases may also have included cells located in the dorsal region of the second auditory field, AII), neurons generally preferred fast FM sweeps. However, a comparison of maps from different animals showed that there was more variability in the distribution of preferred speed responses in the ventral region of the cortex. The directional preference of units for FM sweeps was determined for the sweep speed producing the strongest response. Direction selectivity appeared to be nonrandomly distributed along the dorsoventral axis of AI. In general, units that responded best to upward-directed FM sweeps were located in the more dorsal and ventral aspects of AI while units that responded best to downward-directed FM sweeps were usually located in the mid-region of AI. Direction selectivity was also determined for multiunit responses at each of the three FM sweep speeds. In general, there was a relatively close agreement between the spatial distributions of direction selectivity determined for the strongest response with those calculated for the fast and medium speeds. The spatial distribution of direction selectivity determined for slow FM sweeps deviated somewhat from that determined for the strongest response. Near the dorsoventral center of the mapped areas, the distribution of units that responded best to downward sweeps tended to overlay the distribution of units that responded best to slow speeds, suggesting some spatial covariance of the two parameters.(ABSTRACT TRUNCATED AT 400 WORDS)

Neural selectivity for interaural frequency disparity in cat primary auditory cortex.

Single-unit responses to interaural frequency disparities (IFDs) were examined in 74 neurons in cat primary auditory cortex (AI). Thirty-three of these cells were classified as EE (binaural facilitators), 39 were classified as EI (binaural inhibitors), and 2 were classified as EO (binaural occluders). The best frequency (BF) was presented to the dominant (usually the contralateral) ear while tones of the same or different frequency (either higher or lower than BF) were presented simultaneously to the nondominant (usually the ipsilateral) ear. Most cells displayed sensitivity to IFDs and thus were classified according to the IFD condition that elicited the strongest facilitatory or inhibitory response. The stimulus condition which evoked the strongest binaural response is referred to as the best IFD. For 50 cells (68%), the best IFD response was obtained when tones of different frequency were presented to each ear. Across the entire sample, binaural IFD responses of cortical neurons were categorized into one of three groups: Those preferring a lower frequency than BF in the ipsilateral ear (referred to as the 'lower IFD group'), those preferring a frequency equal to BF (the 'zero IFD group'), or those preferring a frequency higher than BF (the 'higher IFD group'). Among EE cells, approximately one third were maximally facilitated when the ipsilateral ear frequency was lower than BF, one third when it was equal to BF, and one third when it was higher than BF. Among EI cells, 50% exhibited deepest inhibition for higher IFDs with relatively fewer cells showing inhibition for zero or lower IFDs. Overall, EI cells responded over a broader range of IFD conditions than EE cells. Finally, approximately 50% of all units exhibited bimodal responses such that cells classified as EE displayed some inhibitory response characteristics when stimulated with certain IFD conditions and vice versa.

A comparison of monaural and binaural responses to frequency modulated (FM) sweeps in cat primary auditory cortex.

Monaural and binaural single unit responses to frequency-modulated (FM) sweeps were compared in cat primary auditory cortex (AI). Both upward-directed (changing from low to high frequency) and downward-directed (changing from high to low frequency) FM sweeps were presented monaurally and binaurally at five rates of frequency modulation (referred to here as the speed of FM sweep). Two types of binaural FM sweep conditions were presented: (1) like-directed FM sweeps, in which identical FM sweeps were presented to both ears, and (2) opposite-directed FM sweeps, in which one ear was presented with one direction of FM sweep while the other ear was simultaneously presented with the opposite direction of FM sweep. In a sample of 78 cells, 33 cells were classified as EE (binaural facilitatory) and 45 were classified as EI (binaural inhibitory). Ninety-four percent of all units were sensitive to the direction and/or speed of FM sweeps. In general, under binaural stimulus conditions, EE cells responded optimally to like-directed FM sweeps, while EI cells preferred opposite-directed FM sweeps. When tested monaurally, 59% of all cells (both EE and EI) were direction selective, with the majority (76%) preferring downward-directed FM sweeps. When tested binaurally, most direction selective EE cells (60%) preferred upward-directed FM sweeps, while the majority of direction selective EI cells (71%) preferred downward-directed FM sweeps. Our analysis also allowed us to classify inhibitory responses of EI cells as either direction selective (37%) or non-direction selective (63%). For FM speed selectivity under monaural conditions, most EE cells preferred fast FM sweep rates (0.4-0.8 kHz/ms), while approximately equal numbers of EI cells preferred either slow (i.e., 0.05-0.1 kHz/ms) or fast (i.e., 0.4-0.8 kHz/ms) speeds. Under binaural conditions, the majority of EE and EI cells responded best to high speeds when tested with like-directed FM sweeps, while the preferred speed with opposite-directed FM sweeps was more broadly tuned. The results suggest the presence of binaural neural mechanisms underlying cortical FM sweep direction and speed selectivity.

Functional topography of cat primary auditory cortex: representation of tone intensity.

The neuronal response to tones as a function of intensity was topographically studied with multiple-unit recordings in the primary auditory cortex (AI) of barbiturate-anesthetized cats. The spatial distribution of the characteristics of rate/level functions was determined in each of three intensely studied cases and their relationship to the distribution of spectral parameters (sharpness of tuning and responses to broadband transients) in the same animals was determined. The growth of the high-intensity portion of rate/level functions was estimated by linear regression. Locations with monotonically growing high-intensity portions were spatially segregated from locations with nonmonotonic rate/level functions. Two noncontiguous areas with a high degree of nonmonotonicity were observed. One was located at the dorsoventral center of AI, and a second in the dorsal third of AI. The more ventral aggregate of high nonmonotonicity coincided with the region of sharp frequency tuning. The stimulus levels that produced the highest firing rate (strongest response level, SRL) at any sampled location ranged from 10 to 80 dB sound pressure level (SPL). Several spatial aggregates with either high or low SRLs were observed in AI. The region of sharpest tuning was always associated with a region of low SRLs. The response threshold to contralateral tones at the characteristic frequency (CF) ranged from -10 dB SPL to 85 dB SPL with the majority between 0 and 40 dB SPL. The spatial distribution of response thresholds indicated several segregated areas containing clusters with either higher or lower response thresholds. The correlation of response threshold with integrated bandwidth and transient responses was only weak. Low- and high-intensity tones of the same frequency are represented at different locations in AI as judged by the amount of evoked neuronal activity and are largely independent of the frequency organization. The spatial distribution of locations with high monotonicity and low strongest response levels were aligned with the organization of the integrated excitatory bandwidth and covaried with the response strength to broadband stimuli.

Functional topography of cat primary auditory cortex: distribution of integrated excitation.

1. Neuronal responses to tones and transient stimuli were mapped with microelectrodes in the primary auditory cortex (AI) of barbiturate anesthetized cats. Most of the dorsoventral extent of AI was mapped with multiple-unit recordings in the high-frequency domain (between 5.8 and 26.3 kHz) of all six studied cases. The spatial distributions of 1) sharpness of tuning measured with pure tones and 2) response magnitudes to a broadband transient were determined in each of three intensively studied cases. 2. The sharpness of tuning of integrated cluster responses was defined 10 dB above threshold (Q10 dB, integrated excitatory bandwidth). The spatial reconstructions revealed a frequency-independent maximum located near the center of the dorsoventral extent of AI. The sharpness of tuning gradually decreased toward the dorsal and ventral border of AI in all three cases. 3. The sharpness of tuning 40 dB above response threshold was also analyzed (Q40 dB). The Q40 dB values were less than one-half of the corresponding Q10 dB value. The spatial distribution showed a maximum in the center of AI, similar to the Q10 dB distribution. In two out of three cases, restricted additional maxima were recorded dorsal to the main maximum. Overall, Q10 dB and Q40 dB were only moderately correlated, indicating that the integrated excitatory bandwidth at higher stimulus levels can be influenced by additional mechanisms that are not active at lower levels. 4. The magnitude of excitatory responses to a broadband transient (frequency-step response) was determined. The normalized response magnitude varied between less than 1% and up to 100% relative to a characteristic frequency (CF) tone response. The step-response magnitude showed a systematic spatial distribution. An area dorsal to the Q10 dB maximum consistently showed the largest response magnitude surrounded by areas of lower responsivity. A second spatially more restricted maximum was recorded in the ventral-third of each map. Areas with high-transient responsiveness coincided with areas of broad integrated excitatory bandwidth at comparable stimulus levels. 5. The distribution of excitation produced by narrowband and broadband signals suggest that there exists a clear functional organization in the isofrequency domain of AI that is orthogonal to the main cochleotopic organization of the AI. Systematic spatial variations of the integrated excitatory bandwidth reflect underlying cortical processing capacities that may contribute to a parallel analysis of spectral complexity, e.g., spectral shape and contrast, at any given frequency.(ABSTRACT TRUNCATED AT 400 WORDS)

Sensitivity of cat primary auditory cortex (AI) neurons to the direction and rate of frequency modulation.

Responses of 65 single auditory cortex (AI) neurons to frequency-modulated (FM) sweeps with different rates and direction of frequency change were examined quantitatively. Most units responded differentially depending on the characteristics of the FM sweep stimulus. Sixty-five percent of the units encountered responded at least twice as well for one direction of the FM sweep as for the other direction. Of these direction selective neurons, 67% preferred downward-directed FM sweeps (i.e. changing from high to low frequencies) while only 33% preferred upward-directed FM sweeps. The preference for downward-directed FM sweeps was especially clear in EI cells. In addition, cortical neurons often displayed sensitivity to the rate of frequency modulation (speed sensitivity).

Responses of single neurones in cat auditory cortex to time-varying stimuli: frequency-modulated tones of narrow excursion.

In the primary auditory cortex of cats anaesthetized with nitrous oxide, single neurones were examined with respect to their responses to tone bursts and linear modulations of the frequency of an on-going continuous tone. Using FM ramps of 2.0 kHz excursion and varying centre frequency, each of 39 neurones was examined for its preference for the direction of frequency change of a ramp whose centre frequency was varied in and around the neurone's response area. Direction preference was strictly associated with the slopes of the cell's spike count-versus-frequency function over the frequency range covered by the ramp. Preferences for upward- and downward-directed ramps were associated with the low- and high-frequency slopes of the spike count function, respectively. The strength of the cell's direction preference was associated with the relative steepness of the spike count function over the frequency range covered by the ramp. The timing of discharges elicited by the frequency modulations was found to be the sum of the cell's latent period for tone bursts plus the time after ramp onset that the stimulus frequency fell within the neurone's response area. The implications of these data for the processing of narrow and broad frequency-modulated ramps are discussed.