The role of temporal generalization in a temporal discrimination task

Two experiments trained rats to discriminate two or three stimulus durations using a temporal discrimination task. A standard peak shift effect was observed when training was administered with short versus long signals in Experiment 1. Both discrimination accuracy scores and the generalization gradients revealed that shorter intervals were discriminated more accurately, which may be due to the scalar property of timing. In Experiment 2, three signals (short, medium, and long) were associated with three different responses, or two of the intervals were associated with one response (short and long or short and medium) and the other interval with a different response. Here, the short/medium versus long discrimination was learned most readily of the three tasks. The results of both experiments indicated a strong contribution of learning of individual durations combined with scalar generalization gradients, but Experiment 2 indicated that categorical encoding of durations may have also been operating.     

Operant generalization in quail neonates after intradimensional training: Distinguishing positive and negative reinforcement

Operant generalization has been demonstrated in neonates only recently. To investigate the development of intradimensional stimulus control immediately after hatching, northern bobwhite chicks (Colinus virginianus) pecked for brief heat presentations while hearing a high-pitched sound repeated at two constant rates: an S+ tempo signaling a rich reinforcement schedule, alternating with an S− tempo signaling a leaner schedule. Tempo generalization was then assessed in extinction. The expected excitatory gradients were produced after a threshold number of training sessions; unexpectedly, below that threshold, gradients were inhibitory. The chicks’ rapidly developing thermoregulatory capability may have resulted in a change from perceived negative reinforcement initially to positive reinforcement later. Given past research showing excitatory gradients after negative reinforcement, we suggest that these results demonstrate that all negative reinforcement is not equivalent, and, further, that classical conditioning effects require consideration.     

Assessing temporal scales and patterns in time series: Comparing methods based on redundancy analysis

Time-series modelling techniques are powerful tools for studying temporal scaling structures and dynamics present in ecological and other complex systems and are gaining popularity for assessing resilience quantitatively. Among other methods, canonical ordinations based on redundancy analysis are increasingly used for determining temporal scaling patterns that are inherent in ecological data. However, modelling outcomes and thus inference about ecological dynamics and resilience may vary depending on the approaches used. In this study, we compare the statistical performance, logical consistency and information content of two approaches: (i) asymmetric eigenvector maps (AEM) that account for linear trends and (ii) symmetric distance-based Moran's eigenvector maps (MEM), which requires detrending of raw data to remove linear trends prior to analysis. Our comparison is done using long-term water quality data (25 years) from three Swedish lakes. This data set therefore provides the opportunity for assessing how the modelling approach used affects performance and inference in time series modelling. We found that AEM models had consistently more explanatory power than MEM, and in two out of three lakes AEM extracted one more temporal scale than MEM. The scale-specific patterns detected by AEM and MEM were uncorrelated. Also individual water quality variables explaining these patterns differed between methods, suggesting that inferences about systems dynamics are dependent on modelling approach. These findings suggest that AEM might be more suitable for assessing dynamics in time series analysis compared to MEM when temporal trends are relevant. The AEM approach is logically consistent with temporal autocorrelation where earlier conditions can influence later conditions but not vice versa. The symmetric MEM approach, which ignores the asymmetric nature of time, might be suitable for addressing specific questions about the importance of correlations in fluctuation patterns where there are no confounding elements of linear trends or a need to assess causality.     

Time perception and temporal processing levels of the brain

A classification is presented to structure divergent empirical findings on temporal mechanisms of the brain. Proceeding from our time experiences of simultaneity, nonsimultaneity, temporal order, duration, and the subjective present, a classification of thresholds is established that marks distinct processes involved in time perception and the temporal control of movements. On the basis of this classification, evidence has been collected that suggests that perception and action share common timing mechanisms. Furthermore, brain structures involved in temporal aspects of behavior on different time scales have been identified-namely, the cerebellum, the basal ganglia, and circumscribed cortical regions. Another question under debate concerns the representational mode in which time is represented in the brain; models suggest neuronal oscillations or interval-based mechanisms.     

How training and testing histories affect generalization: a test of simple neural networks

We show that a simple network model of associative learning can reproduce three findings that arise from particular training and testing procedures in generalization experiments: the effect of (i) 'errorless learning', (ii) extinction testing on peak shift, and (iii) the central tendency effect. These findings provide a true test of the network model which was developed to account for other phenomena, and highlight the potential of neural networks to study the phenomena that depend on sequences of experiences with many stimuli. Our results suggest that at least some such phenomena, e.g. stimulus range effects, may derive from basic mechanisms of associative memory rather than from more complex memory processes.     

Relative temporal representations in Pavlovian conditioning

The transfer of relative temporal representations was assessed in a series of three experiments. In each experiment, rats (Rattus norvegicus) received one set of conditioned stimulus (CS) and intertrial interval (ITI) durations in Phase 1 and another set in Phase 2. The ratio between the CS and ITI intervals was either changed or maintained across phases. On the hypothesis that relative temporal representations are learned, groups receiving maintained temporal ratios across phases were expected to display greater change in responding upon encountering the new intervals. When the CS duration decreased across phases, maintaining the temporal ratio did lead to greater change in Day 1 of Phase 2 towards the final pattern of responding. However, when the CS increased across phases, maintaining the temporal ratio across phases did not facilitate adjustment to the new intervals, suggesting that extinction of previously reinforced times induced new learning. These results provide evidence that under some conditions, relative relationships in temporal maps may survive transformation-of-scale, like relative relationships in spatial maps.     

Electrophysiological and psychophysical quantification of temporal summation in the human nociceptive system

Animal experiments have shown that the nociceptive reflex can be used as an indicator of central temporal integration in the nociceptive system. The aim of the present study on humans was to investigate whether the nociceptive reflex, evoked by repetitive strong electrical sural nerve stimuli, increased when summation was reported by the volunteers. The reflexes were recorded from the biceps femoris and rectus femoris muscles in eight volunteers following a series of stimulations at 0.1, 1, 2, and 3 Hz. Each series consisted of five consecutive stimuli. Using 0.1- and 1-Hz stimulation, the reflex was not facilitated in the course of the five consecutive stimuli. Following 2- and 3-Hz stimulation, the reflex size (root mean square amplitude) increased significantly during the course of the fifth stimulus. This reflex facilitation was followed by a significant increase (summation) in the pain magnitude when compared with 1- and 0.1-Hz stimulation. Furthermore, the threshold for psychophysical summation could be determined. This threshold (stimulus intensity) decreased when the stimulus frequency (1–5 Hz) of the five consecutive stimuli was increased. The nociceptive reflex and the psychophysical summation threshold might be used to clarify and quantify aspects of temporal summation within the human nociceptive system.     

生物的时间地带性及其农学涵义

生物的时间地带性可表述为生物有机体为谋求生存阈限的扩展,突破地理地带性（暂称为空间地带性）的局限,对地理地带性加以剪裁,连缀,排除不适宜自身生存的环境时段,截取适宜自身生存的环境时段所构建的生存时空体;生物的时间地带性是生物对地球运动（自转与公转）造成的节律适应的结果,时间地带性依赖于空间地带性,不同的空间地带内,时间带谱的基带不同,并在空间地带性的基础上呈现为螺旋状往复循环,是生态要素的时间周期性变化规律的体现;时间地带性与空间地带性一样,都是依赖于尺度（时间尺度与空间尺工）的生态要素分布过程,带谱的成分,范围均随着尺度的变化而变化,生物通过不同的生态策略来适应这种变化;人类农业活动已经取得一些成效,但只能在小尺度上调节时间地带性,不能改变空间地带性。     

Spatial and temporal stochastic interaction in neuronal assemblies

Summary The observation of various types of spatio-temporal correlations in spike-patterns of multiple cortical neurons has shifted attention from rate coding paradigms to computational processes based on the precise timing of spikes in neuronal ensembles. In the present work we develop the notion of “spatial” and “temporal interaction” which provides measures for statistical dependences in coupled stochastic processes like multiple unit spike trains. We show that the classical Willshaw network and Abeles’ synfire chain model both reveal a moderate spatial interaction, but only the synfire chain model reveals a positive temporal interaction, too. Systems that maximize temporal interaction are shown to be almost deterministic globally, but posses almost unpredictable firing behavior on the single unit level.     

Temporal generalization accounts for response resurgence in the peak procedure

The peak interval (PI) procedure is commonly used to evaluate animals' ability to produce timed intervals. It consists of presenting fixed interval (FI) schedules in which some of the trials are replaced by extended non-reinforced trials. Responding will often resume (resurge) at the end of the non-reinforced trials unless precautions are taken to prevent it. Response resurgence was replicated in rats and pigeons. Variation of the durations of the FI and the non-reinforced probe trials showed it to be dependent on the time when reinforcement is expected. Timing of both the normal time to reinforcement, and the subsequent time to reinforcement during the probe trials followed Weber's law. A quantitative model of resurgence is described, suggesting how animals respond to the signaling properties of reinforcement omission. Model results were simulated using a stochastic binary counter.     

Model based generalization analysis of common spatial pattern in brain computer interfaces

In the motor imagery based Brain Computer Interface (BCI) research, Common Spatial Pattern (CSP) algorithm is used widely as a spatial filter on multi-channel electroencephalogram (EEG) recordings. Recently the overfitting effect of CSP has been gradually noticed, but what influence the overfitting is still unclear. In this work, the generalization of CSP is investigated by a simple linear mixing model. Several factors in this model are discussed, and the simulation results indicate that channel numbers and the correlation between signals influence the generalization of CSP significantly. A larger number of training trials and a longer time length of the trial would prevent overfitting. The experiments on real data also verify our conclusion.     

Pollinator diversity affects plant reproduction and recruitment: the tradeoffs of generalization

One outstanding and unsolved challenge in ecology and conservation biology is to understand how pollinator diversity affects plant performance. Here, we provide evidence of the functional role of pollination diversity in a plant species, Erysimum mediohispanicum (Brassicaceae). Pollinator abundance, richness and diversity as well as plant reproduction and recruitment were determined in eight plant populations. We found that E. mediohispanicum was generalized both at the regional and local (population) scale, since its flowers were visited by more than 100 species of insects with very different morphology, size and behaviour. However, populations differed in the degree of generalization. Generalization correlated with pollinator abundance and plant population size, but not with habitat, ungulate damage intensity, altitude or spatial location. More importantly, the degree of generalization had significant consequences for plant reproduction and recruitment. Plants from populations with intermediate generalization produced more seeds than plants from populations with low or high degrees of generalization. These differences were not the result of differences in number of flowers produced per plant. In addition, seedling emergence in a common garden was highest in plants from populations with intermediate degree of generalization. This outcome suggests the existence of an optimal level of generalizations even for generalized plant species.     

Species temporal variability: epigaeic ant assemblages and management for abundance and scarcity

Summary Ant assemblages in five closely-spaced point habitats were studied over a three-year period in a subtropical area of South Africa. A total of 20,746 individuals in 56 species were sampled with pitfall traps. Rapid species turnover occurred during short periods after longer intervals of relatively constant species composition. There was annual cycling of total numbers of individuals. The temporal variance for total number of individuals within each habitat increased significantly with mean population size by a logarithmic regression. This indicates that population fluctuations are dependent upon the mean, with a distinctive relationship. The total number of species was cyclical, with annual and approximately half-yearly cycles. Each habitat nevertheless carried a characteristic mean number of species. The temporal variance of species richness showed a highly significant positive logarithmic relationship with the mean number of species. Pheidole megacephala (F.) accounted for 45% of all individuals. Its overall population showed annual cycles. Ph. megacephala population trend was the main determinant for seasonal variation of the whole assemblage in most habitats. Percentage dominance was positively linearly correlated with the variance and with mean population level. Although Ph. megacephala population variance increased with the mean, proportionate variability did not, indicating that high populations are no more variable than low populations. Five variability measures were compared with each other on ant assemblages containing only the common species (1st quartile) and then including the increasingly rare species (1st and 2nd, 1st, 2nd and 3rd, and all quartiles). Williamson's Variability Measure, using logarithmic or inverse sine transformations, gave variable results depending whether or not the rarer species were included in the analysis. The Coefficient of Fluctuation also gave variable results depending on assemblage size. The Coefficient of Variation and Proportionate Variability Measure gave similar and consistent results, indicating that common species are not more variable than rare ones. The rarer species are stenotopic and tend always to be rare, whereas the common species are eurytopic, highly competitive and demographically volatile. Control measures for common ants, and conservation of rare ants, should take cognizance of these trends.     

Inverted behavioural responses in wild-type Rhodobacter sphaeroides to temporal stimuli

Both aerobically and photosynthetically grown wild-type Rhodobacter sphaeroides swarmed through soft nutrient agar. However, individual aerobically and photosynthetically grown tethered cells showed different responses to steps in concentrations of some attractants. Photosynthetically grown cells showed little response to a step-up in attractant, but large response to a step-down. Aerobically grown cells showed a large but opposite response to a step-up of chemoeffectors such as succinate and aspartate. The responses in che operon deletion mutants were also investigated and indicated that the aerobic response may depend on the protein products of che operon 1.     

Impact of microphytobenthos and macroinfauna on temporal variation of benthic metabolism in shallow coastal sediments

The impact of microphytobenthos and different abundances of macrofauna (Nereis diversicolor) on temporal variation of benthic metabolism was investigated in laboratory microcosms. Measurements primarily included diurnal fluxes of O₂ and CO₂ as well as sediment profiles of Chlorophyll a and extracellular polymeric substances (EPS). Net and gross primary production (2-5 and 4-7 mmol CO₂ m^{− 2} h^{− 1}, respectively) were relatively stable in both defaunated and faunated sediment throughout a 12 h light period. The CO₂ release from sediments immediately after onset of darkness ranged from 1.5 to 3.5 mmol CO₂ m^{− 2} h^{− 1} followed by a consistent decrease during the next 12 h in the dark. The decrease was more conspicuous in faunated (about 50%) than defaunated (9%) sediment. Total carbon oxidation was in both cases fuelled primarily by microphytobenthic biomass, while EPS only contributed by 1-4%. Diurnal measurements of Nereis diversicolor ventilation activity showed a significant decrease in the dark that corresponds well to the observed decrease in total metabolic activity. It is concluded that changes in solute exchange associated with animals and burrows (e.g. microbial respiration) is a major controlling factor for total sediment metabolism. In general, the faunal impact was evident as about 50% enhanced CO₂ release in the dark, while net primary production was reduced by 30-50%. The turnover time of produced organic carbon is therefore considerably shorter in the presence than absence of macrofauna. Thus, the daily average exchange of CO₂ was almost balanced in bioturbated sediment with a 43% share of carbon oxidation mediated by direct faunal respiration. Defaunated sediment was net autotrophic with daily primary production exceeding microbial carbon oxidation by 40%. The present study clearly demonstrates that knowledge on interactions between microphytobenthos and macrofauna is essential for understanding carbon dynamics in shallow sediments.     

Daily temporal organization of laying in Japanese quail: variability and heritability

In birds, many behavioral and physiological processes that occur during reproduction show daily rhythms in response to environmental temporal constraints. In this study, the individual daily organization of laying and its genetic determinant in Japanese quail (Coturnix coturnix japonica) were analyzed. For this purpose, the oviposition time of 102 randomly chosen females, maintained in long-day photoperiodic conditions (LD 14h:10h) for 1 mon was observed and extreme phenotypes selected. Laying is characterized by two parameters: oviposition interval and laying hour.

The birds showed a specific time of laying during 24 h. All eggs were laid in the afternoon between 6.5 and 14 h after lights on (HALO). Two laying profiles were determined: 20% of females with an oviposition interval greater than 24 h (24.7±0.2 h) (the “delayed” profile) laid progressively later each day until a pause day. The remaining 80% of the females laid at the same time each day, with few pause days and an oviposition interval close to 24 h (24.0±0.2 h) (the “stable” profile). Among the females, showing this last profile, an intra-individual stability and an inter-individual variability of laying hour was established. Two extreme laying phenotypes were then determined: the “early” phenotype (E) for females laying on average between 7.5 and 9.5 HALO and the “late” phenotype (L) for females laying between 12.5 and 14 HALO.

In order to study the genetic basis of the laying hour, three females of each extreme phenotype were selected and crossed with two different males. The E and L females produced 57 F1E and 42 F1L daughters, respectively. F1 females displayed both laying profiles. However, the proportion of females displaying a “delayed” profile was higher in the L line (50%) than in the E line (29.8%). For the “stable” daughters, artificial selection induced an advance in laying hour of 4.7% for the E line and a delay of 4.7% for the L line. Realized heritability was estimated at 0.5. Moreover, the laying hour of the daughters was correlated positively to that of the mothers (N=61;r=0.45). These results support the notion of heritability of oviposition time in Japanese quail.     

Chemosensory processing in the fruit fly,Drosophila melanogaster: generalization of a feeding response reveals overlapping odour representations

Insects are capable of detecting, and discriminating between, a very large number of odours. The biological relevance of many of those odours, particularly those related to food, must first be learned. Given that the number of sensory receptors and antennal lobe (AL) glomeruli is limited relative to the number of odours that must be detectable, this ability implies that the olfactory system makes use of a combinatorial coding scheme whereby each sensory cell or AL projection neuron can participate in coding for several different odours. An important step in understanding this coding scheme is to behaviourally quantify the degree to which sets of odours are discriminable. Here we evaluate odour discriminability in the fruit fly,Drosophila melanogaster, by first conditioning individual flies to not respond to any of several odourants using a nonassociative conditioning protocol (habituation). We show that flies habituate unconditioned leg movement responses to both mechanosensory and olfactory stimulation over 25 unreinforceed trials. Habituation is retained for at least 2 h and is subject to dishabituation. Finally, we test the degree to which the conditioned response generalizes to other odourants based on molecular features of the odourants (e.g. carbon chain length and the presence of a target functional group). These tests reveal predictable generalization gradients across these molecular features. These data substantiate the claim that these features are relevant coding dimensions in the fruit fly olfactory system, as has been shown for other insect and vertebrate species.     

Daily temporal organization of laying in Japanese quail: variability and heritability

In birds, many behavioral and physiological processes that occur during reproduction show daily rhythms in response to environmental temporal constraints. In this study, the individual daily organization of laying and its genetic determinant in Japanese quail (Coturnix coturnix japonica) were analyzed. For this purpose, the oviposition time of 102 randomly chosen females, maintained in long-day photoperiodic conditions (LD 14h:10h) for 1 mon was observed and extreme phenotypes selected. Laying is characterized by two parameters: oviposition interval and laying hour.

The birds showed a specific time of laying during 24 h. All eggs were laid in the afternoon between 6.5 and 14 h after lights on (HALO). Two laying profiles were determined: 20% of females with an oviposition interval greater than 24 h (24.7±0.2 h) (the “delayed” profile) laid progressively later each day until a pause day. The remaining 80% of the females laid at the same time each day, with few pause days and an oviposition interval close to 24 h (24.0±0.2 h) (the “stable” profile). Among the females, showing this last profile, an intra-individual stability and an inter-individual variability of laying hour was established. Two extreme laying phenotypes were then determined: the “early” phenotype (E) for females laying on average between 7.5 and 9.5 HALO and the “late” phenotype (L) for females laying between 12.5 and 14 HALO.

In order to study the genetic basis of the laying hour, three females of each extreme phenotype were selected and crossed with two different males. The E and L females produced 57 F1E and 42 F1L daughters, respectively. F1 females displayed both laying profiles. However, the proportion of females displaying a “delayed” profile was higher in the L line (50%) than in the E line (29.8%). For the “stable” daughters, artificial selection induced an advance in laying hour of 4.7% for the E line and a delay of 4.7% for the L line. Realized heritability was estimated at 0.5. Moreover, the laying hour of the daughters was correlated positively to that of the mothers (N=61;r=0.45). These results support the notion of heritability of oviposition time in Japanese quail.