Properties of auditory stream formation

A sequence of sounds may be heard as coming from a single source (called fusion or coherence) or from two or more sources (called fission or stream segregation). Each perceived source is called a 'stream'. When the differences between successive sounds are very large, fission nearly always occurs, whereas when the differences are very small, fusion nearly always occurs. When the differences are intermediate in size, the percept often 'flips' between one stream and multiple streams, a property called 'bistability'. The flips do not generally occur regularly in time. The tendency to hear two streams builds up over time, but can be partially or completely reset by a sudden change in the properties of the sequence or by switches in attention. Stream formation depends partly on the extent to which successive sounds excite different 'channels' in the peripheral auditory system. However, other factors can play a strong role; multiple streams may be heard when successive sounds are presented to the same ear and have essentially identical excitation patterns in the cochlea. Differences between successive sounds in temporal envelope, fundamental frequency, phase spectrum and lateralization can all induce a percept of multiple streams. Regularities in the temporal pattern of elements within a stream can help in stabilizing that stream.     

Auditory Streaming as an Online Classification Process with Evidence Accumulation

When human subjects hear a sequence of two alternating pure tones, they often perceive it in one of two ways: as one integrated sequence (a single "stream" consisting of the two tones), or as two segregated sequences, one sequence of low tones perceived separately from another sequence of high tones (two "streams"). Perception of this stimulus is thus bistable. Moreover, subjects report on-going switching between the two percepts: unless the frequency separation is large, initial perception tends to be of integration, followed by toggling between integration and segregation phases. The process of stream formation is loosely named “auditory streaming”. Auditory streaming is believed to be a manifestation of human ability to analyze an auditory scene, i.e. to attribute portions of the incoming sound sequence to distinct sound generating entities. Previous studies suggested that the durations of the successive integration and segregation phases are statistically independent. This independence plays an important role in current models of bistability. Contrary to this, we show here, by analyzing a large set of data, that subsequent phase durations are positively correlated. To account together for bistability and positive correlation between subsequent durations, we suggest that streaming is a consequence of an evidence accumulation process. Evidence for segregation is accumulated during the integration phase and vice versa; a switch to the opposite percept occurs stochastically based on this evidence. During a long phase, a large amount of evidence for the opposite percept is accumulated, resulting in a long subsequent phase. In contrast, a short phase is followed by another short phase. We implement these concepts using a probabilistic model that shows both bistability and correlations similar to those observed experimentally.     

Perceptual organization of sound begins in the auditory periphery

Segmenting the complex acoustic mixture that makes a typical auditory scene into relevant perceptual objects is one of the main challenges of the auditory system [1], for both human and nonhuman species. Several recent studies indicate that perceptual auditory object formation, or "streaming," may be based on neural activity within the auditory cortex and beyond [2, 3]. Here, we find that scene analysis starts much earlier in the auditory pathways. Single units were recorded from a peripheral structure of the mammalian auditory brainstem, the cochlear nucleus. Peripheral responses were similar to cortical responses and displayed all of the functional properties required for streaming, including multisecond adaptation. Behavioral streaming was also measured in human listeners. Neurometric functions derived from the peripheral responses predicted accurately behavioral streaming. This reveals that subcortical structures may already contribute to the analysis of auditory scenes. This finding is consistent with the observation that species lacking a neocortex can still achieve and benefit from behavioral streaming [4]. For humans, we argue that auditory scene analysis of complex scenes is probably based on interactions between subcortical and cortical neural processes, with the relative contribution of each stage depending on the nature of the acoustic cues forming the streams.     

Can two streams of auditory information be processed simultaneously? Evidence from the gleaning bat Antrozous pallidus

A tenet of auditory scene analysis is that we can fully process only one stream of auditory information at a time. We tested this assumption in a gleaning bat, the pallid bat (Antrozous pallidus) because this bat uses echolocation for general orientation, and relies heavily on prey-generated sounds to detect and locate its prey. It may therefore encounter situations in which the echolocation and passive listening streams temporally overlap. Pallid bats were trained to a dual task in which they had to negotiate a wire array, using echolocation, and land on one of 15 speakers emitting a brief noise burst in order to obtain a food reward. They were forced to process both streams within a narrow 300 to 500 ms time window by having the noise burst triggered by the bats initial echolocation pulses as it approached the wire array. Relative to single task controls, echolocation and passive sound localization performance was slightly, but significantly, degraded. The bats also increased echolocation interpulse intervals during the dual task, as though attempting to reduce temporal overlap between the signals. These results suggest that the bats, like humans, have difficulty in processing more than one stream of information at a time.     

Audio-visual speech timing sensitivity is enhanced in cluttered conditions

Events encoded in separate sensory modalities, such as audition and vision, can seem to be synchronous across a relatively broad range of physical timing differences. This may suggest that the precision of audio-visual timing judgments is inherently poor. Here we show that this is not necessarily true. We contrast timing sensitivity for isolated streams of audio and visual speech, and for streams of audio and visual speech accompanied by additional, temporally offset, visual speech streams. We find that the precision with which synchronous streams of audio and visual speech are identified is enhanced by the presence of additional streams of asynchronous visual speech. Our data suggest that timing perception is shaped by selective grouping processes, which can result in enhanced precision in temporally cluttered environments. The imprecision suggested by previous studies might therefore be a consequence of examining isolated pairs of audio and visual events. We argue that when an isolated pair of cross-modal events is presented, they tend to group perceptually and to seem synchronous as a consequence. We have revealed greater precision by providing multiple visual signals, possibly allowing a single auditory speech stream to group selectively with the most synchronous visual candidate. The grouping processes we have identified might be important in daily life, such as when we attempt to follow a conversation in a crowded room.     

Neurophysiological mechanisms of auditory adaptation. II. Poststimulus effects

The problem of interaction of spike neuronal activity evoked by successive sounds in single elements of auditory system is considered. The forward masking situation, when pairs of signals are presented independently, as well as the condition of long sequences of signals with different on-of ratios are analyzed. The strong increase of a diversity of single units ability to reproduce fast sequences really observed from the lowest to the higher nuclei of an auditory pathway. Complex units, reacting only on "new" signals, appear from midbrain region of auditory pathway. However such elements are found out usually not in a direct lemniscal auditory way, but in surrounding nuclei. While poststimulus adaptation to specified type of signals usually causes the considerable increase in threshold of detection, differential sensitivity to small changes can remain quite high. This aspect of auditory sensation remains poorly investigated both in physiological and in psychophysical experiments.     

Functional architecture of auditory cortex

Three complementary approaches demonstrate new types of organization in rodent, feline and primate auditory cortex, as well as differences in processing between auditory and visual cortex. First, connectional work reveals patterns of thalamocortical and corticocortical input unique to the auditory cortex. Second, physiological studies find multiple, interleaved auditory processing modules related to corticocortical connections and embedded in the isofrequency gradient. Third, functional analyses demonstrate independent processing streams for sound localization and identification analogous to the 'what' and 'where' streams in visual cortex, although the modular arrangements are modality-specific. Taken together, these data show that the auditory cortex has common and unique functional substrates.     

Segregation and Integration of Auditory Streams when Listening to Multi-Part Music

In our daily lives, auditory stream segregation allows us to differentiate concurrent sound sources and to make sense of the scene we are experiencing. However, a combination of segregation and the concurrent integration of auditory streams is necessary in order to analyze the relationship between streams and thus perceive a coherent auditory scene. The present functional magnetic resonance imaging study investigates the relative role and neural underpinnings of these listening strategies in multi-part musical stimuli. We compare a real human performance of a piano duet and a synthetic stimulus of the same duet in a prioritized integrative attention paradigm that required the simultaneous segregation and integration of auditory streams. In so doing, we manipulate the degree to which the attended part of the duet led either structurally (attend melody vs. attend accompaniment) or temporally (asynchronies vs. no asynchronies between parts), and thus the relative contributions of integration and segregation used to make an assessment of the leader-follower relationship. We show that perceptually the relationship between parts is biased towards the conventional structural hierarchy in western music in which the melody generally dominates (leads) the accompaniment. Moreover, the assessment varies as a function of both cognitive load, as shown through difficulty ratings and the interaction of the temporal and the structural relationship factors. Neurally, we see that the temporal relationship between parts, as one important cue for stream segregation, revealed distinct neural activity in the planum temporale. By contrast, integration used when listening to both the temporally separated performance stimulus and the temporally fused synthetic stimulus resulted in activation of the intraparietal sulcus. These results support the hypothesis that the planum temporale and IPS are key structures underlying the mechanisms of segregation and integration of auditory streams, respectively.     

Neural Processing of Short-Term Recurrence in Songbird Vocal Communication

Background

Many situations involving animal communication are dominated by recurring, stereotyped signals. How do receivers optimally distinguish between frequently recurring signals and novel ones? Cortical auditory systems are known to be pre-attentively sensitive to short-term delivery statistics of artificial stimuli, but it is unknown if this phenomenon extends to the level of behaviorally relevant delivery patterns, such as those used during communication.

Methodology/Principal Findings

We recorded and analyzed complete auditory scenes of spontaneously communicating zebra finch (Taeniopygia guttata) pairs over a week-long period, and show that they can produce tens of thousands of short-range contact calls per day. Individual calls recur at time scales (median interval 1.5 s) matching those at which mammalian sensory systems are sensitive to recent stimulus history. Next, we presented to anesthetized birds sequences of frequently recurring calls interspersed with rare ones, and recorded, in parallel, action and local field potential responses in the medio-caudal auditory forebrain at 32 unique sites. Variation in call recurrence rate over natural ranges leads to widespread and significant modulation in strength of neural responses. Such modulation is highly call-specific in secondary auditory areas, but not in the main thalamo-recipient, primary auditory area.

Conclusions/Significance

Our results support the hypothesis that pre-attentive neural sensitivity to short-term stimulus recurrence is involved in the analysis of auditory scenes at the level of delivery patterns of meaningful sounds. This may enable birds to efficiently and automatically distinguish frequently recurring vocalizations from other events in their auditory scene.     

Directed attention eliminates 'change deafness' in complex auditory scenes

In natural environments that contain multiple sound sources, acoustic energy arising from the different sources sums to produce a single complex waveform at each of the listener's ears. The auditory system must segregate this waveform into distinct streams to permit identification of the objects from which the signals emanate [1]. Although the processes involved in stream segregation are now reasonably well understood [1, 2 and 3], little is known about the nature of our perception of complex auditory scenes. Here, we examined complex scene perception by having listeners detect a discrete change to an auditory scene comprising multiple concurrent naturalistic sounds. We found that listeners were remarkably poor at detecting the disappearance of an individual auditory object when listening to scenes containing more than four objects, but they performed near perfectly when their attention was directed to the identity of a potential change. In the absence of directed attention, this "change deafness" [4] was greater for objects arising from a common location in space than for objects separated in azimuth. Change deafness was also observed for changes in object location, suggesting that it may reflect a general effect of the dependence of human auditory perception on attention.     

Interaction of Streaming and Attention in Human Auditory Cortex

Serially presented tones are sometimes segregated into two perceptually distinct streams. An ongoing debate is whether this basic streaming phenomenon reflects automatic processes or requires attention focused to the stimuli. Here, we examined the influence of focused attention on streaming-related activity in human auditory cortex using magnetoencephalography (MEG). Listeners were presented with a dichotic paradigm in which left-ear stimuli consisted of canonical streaming stimuli (ABA_ or ABAA) and right-ear stimuli consisted of a classical oddball paradigm. In phase one, listeners were instructed to attend the right-ear oddball sequence and detect rare deviants. In phase two, they were instructed to attend the left ear streaming stimulus and report whether they heard one or two streams. The frequency difference (ΔF) of the sequences was set such that the smallest and largest ΔF conditions generally induced one- and two-stream percepts, respectively. Two intermediate ΔF conditions were chosen to elicit bistable percepts (i.e., either one or two streams). Attention enhanced the peak-to-peak amplitude of the P1-N1 complex, but only for ambiguous ΔF conditions, consistent with the notion that automatic mechanisms for streaming tightly interact with attention and that the latter is of particular importance for ambiguous sound sequences.     

On the cyclic nature of perception in vision versus audition

Does our perceptual awareness consist of a continuous stream, or a discrete sequence of perceptual cycles, possibly associated with the rhythmic structure of brain activity? This has been a long-standing question in neuroscience. We review recent psychophysical and electrophysiological studies indicating that part of our visual awareness proceeds in approximately 7–13 Hz cycles rather than continuously. On the other hand, experimental attempts at applying similar tools to demonstrate the discreteness of auditory awareness have been largely unsuccessful. We argue and demonstrate experimentally that visual and auditory perception are not equally affected by temporal subsampling of their respective input streams: video sequences remain intelligible at sampling rates of two to three frames per second, whereas audio inputs lose their fine temporal structure, and thus all significance, below 20–30 samples per second. This does not mean, however, that our auditory perception must proceed continuously. Instead, we propose that audition could still involve perceptual cycles, but the periodic sampling should happen only after the stage of auditory feature extraction. In addition, although visual perceptual cycles can follow one another at a spontaneous pace largely independent of the visual input, auditory cycles may need to sample the input stream more flexibly, by adapting to the temporal structure of the auditory inputs.     

A model for intramolecular recombination of herpesvirus DNA leading to the formation of circular, circular-linear and fragmented genomes.

A model is presented for intramolecular recombination of herpesvirus DNA. It is proposed that the terminal repeat sequences of the viral DNA contain insertion sequences which may integrate with homologous repeat sequences between the long (L) and short (S) components. In class 2 herpes-virus DNA (as defined by Honess &; Watson, 1977) in which the repeat sequences flank the S component only, circular-linear DNA molecules can be formed as an intermediate step. Reorientation of the S component leads to the formation of two DNA isomers. In class 3 herpesvirus DNA in which repeat sequences flank both the L and S components, either circular-linear or 8-shaped DNA molecules are proposed as intermediates leading to the formation of four DNA isomers. Fragmentation of the S component could lead to the formation of small circular DNA molecules.     

Functional brain networks underlying perceptual switching: auditory streaming and verbal transformations

Recent studies have shown that auditory scene analysis involves distributed neural sites below, in, and beyond the auditory cortex (AC). However, it remains unclear what role each site plays and how they interact in the formation and selection of auditory percepts. We addressed this issue through perceptual multistability phenomena, namely, spontaneous perceptual switching in auditory streaming (AS) for a sequence of repeated triplet tones, and perceptual changes for a repeated word, known as verbal transformations (VTs). An event-related fMRI analysis revealed brain activity timelocked to perceptual switching in the cerebellum for AS, in frontal areas for VT, and the AC and thalamus for both. The results suggest that motor-based prediction, produced by neural networks outside the auditory system, plays essential roles in the segmentation of acoustic sequences both in AS and VT. The frequency of perceptual switching was determined by a balance between the activation of two sites, which are proposed to be involved in exploring novel perceptual organization and stabilizing current perceptual organization. The effect of the gene polymorphism of catechol-O-methyltransferase (COMT) on individual variations in switching frequency suggests that the balance of exploration and stabilization is modulated by catecholamines such as dopamine and noradrenalin. These mechanisms would support the noteworthy flexibility of auditory scene analysis.     

Stream noise,habitat filtering,and the phenotypic and phylogenetic structure of Neotropical anuran assemblages

The structure of assemblages may be determined by interspecific interactions or environmental factors (e.g. competition and habitat filtering). Since communication between conspecific and heterospecific affects fitness of individuals, habitat characteristics that prevent communication could determine habitat use and co-occurrence of species. However, at present there are few studies, most with birds, testing the relationship between sensory ecology and community ecology. Abiotic noise on streams could impede the detection and decoding of auditory signals by receivers through a process named auditory masking. Therefore, we tested the role of abiotic noise on streams as a habitat characteristic influencing the phenotypic and phylogenetic structure of Neotropical anuran assemblages. We tested this hypothesis using data of male body size, call frequency, calling place (alongside and away from streams), and phylogenetic relationship of 110 and 38 anuran species at regional and local scale, respectively. After we found quantitative evidence suggesting that call frequency and body size are conserved phenotypic traits, we found that assemblages alongside streams exhibit both phenotypic and phylogenetic clustering, while assemblages away from streams exhibit both phenotypic and phylogenetic overdispersion. These results offer quantitative evidence suggesting a role of noise on streams promoting a process of habitat filtering and affecting the structure of anuran assemblages alongside streams both at Neotropical and local scale. This is the first study using modern phylogenetic comparative metrics for covering potential causes of phenotypic and phylogenetic structure of anuran assemblages, and one of the few testing a link between community ecology and the evolutionary biology of acoustic communication to understand the processes mediating species co-occurrence in vertebrates.     

Reliability and representational bandwidth in the auditory cortex

It is unclear why there are so many more neurons in sensory cortex than in the sensory periphery. One possibility is that these "extra" neurons are used to overcome cortical noise and faithfully represent the acoustic stimulus. Another possibility is that even after overcoming cortical noise, there is "excess representational bandwidth" available and that this bandwidth is used to represent conjunctions of auditory and nonauditory information for computation. Here, we discuss recent data about neuronal reliability in auditory cortex showing that cortical noise may not be as high as was previously believed. Although at present, the data suggest that auditory cortex neurons can be more reliable than those in the visual cortex, we speculate that the principles governing cortical computation are universal and that visual and other cortical areas can also exploit strategies based on similarly high-fidelity activity.     

The riparian community of north-west European lowland streams

• 1 Riparian plant communities along lowland streams have been affected by man for the past 15000 years. During this period climatological changes have also taken place. The present situation in north-west Europe is a result of these influences so that the choice of reference situations is necessarily arbitrary.
• 2 Dominant abiotic factors that influence riparian vegetation are local hydrology and soil composition, and the pH, lime content and nutrient concentration of the feeding ground water.
• 3 Riparian plant communities are described as phytosociological units. Depending on abiotic characteristics, sequences from spring to lower course and from the stream bank to the boundaries of the stream valley are reported. The Pruno-Fraxinetum association is, and has been for thousands of years, one of the main riparian vegetation types of (lowland) streams.
• 4 In many European countries, restoration projects are being performed or planned. One of the restoration measures is the planting of trees or shrubs along streams as buffer strips. A phytosociological approach, based on knowledge of the local hydrology and nutrient/ion availability, is better by far than the general practice of standard plantation. Such an approach requires research in advance, which increases the initial costs of projects, but can be expected to provide far better results in the long term, and ultimately save money.

     

A neurocomputational model of stimulus-specific adaptation to oddball and Markov sequences

Stimulus-specific adaptation (SSA) occurs when the spike rate of a neuron decreases with repetitions of the same stimulus, but recovers when a different stimulus is presented. It has been suggested that SSA in single auditory neurons may provide information to change detection mechanisms evident at other scales (e.g., mismatch negativity in the event related potential), and participate in the control of attention and the formation of auditory streams. This article presents a spiking-neuron model that accounts for SSA in terms of the convergence of depressing synapses that convey feature-specific inputs. The model is anatomically plausible, comprising just a few homogeneously connected populations, and does not require organised feature maps. The model is calibrated to match the SSA measured in the cortex of the awake rat, as reported in one study. The effect of frequency separation, deviant probability, repetition rate and duration upon SSA are investigated. With the same parameter set, the model generates responses consistent with a wide range of published data obtained in other auditory regions using other stimulus configurations, such as block, sequential and random stimuli. A new stimulus paradigm is introduced, which generalises the oddball concept to Markov chains, allowing the experimenter to vary the tone probabilities and the rate of switching independently. The model predicts greater SSA for higher rates of switching. Finally, the issue of whether rarity or novelty elicits SSA is addressed by comparing the responses of the model to deviants in the context of a sequence of a single standard or many standards. The results support the view that synaptic adaptation alone can explain almost all aspects of SSA reported to date, including its purported novelty component, and that non-trivial networks of depressing synapses can intensify this novelty response.     

Radial and spiral stream formation in Proteus mirabilis colonies

The enteric bacterium Proteus mirabilis, which is a pathogen that forms biofilms in vivo, can swarm over hard surfaces and form a variety of spatial patterns in colonies. Colony formation involves two distinct cell types: swarmer cells that dominate near the surface and the leading edge, and swimmer cells that prefer a less viscous medium, but the mechanisms underlying pattern formation are not understood. New experimental investigations reported here show that swimmer cells in the center of the colony stream inward toward the inoculation site and in the process form many complex patterns, including radial and spiral streams, in addition to previously-reported concentric rings. These new observations suggest that swimmers are motile and that indirect interactions between them are essential in the pattern formation. To explain these observations we develop a hybrid model comprising cell-based and continuum components that incorporates a chemotactic response of swimmers to a chemical they produce. The model predicts that formation of radial streams can be explained as the modulation of the local attractant concentration by the cells, and that the chirality of the spiral streams results from a swimming bias of the cells near the surface of the substrate. The spatial patterns generated from the model are in qualitative agreement with the experimental observations.     

Effects of habitat features on size-biased predation on salmon by bears

Predators can drive trait divergence among populations of prey by imposing differential selection on prey traits. Habitat characteristics can mediate predator selectivity by providing refuge for prey. We quantified the effects of stream characteristics on biases in the sizes of spawning salmon caught by bears (Ursus arctos and U. americanus) on the central coast of British Columbia, Canada by measuring size-biased predation on spawning chum (Oncorhynchus keta) and pink (O. gorbuscha) salmon in 12 streams with varying habitat characteristics. We tested the hypotheses that bears would catch larger than average salmon (size-biased predation) and that this bias toward larger fish would be higher in streams that provide less protection to spawning salmon from predation (e.g., less pools, wood, undercut banks). We then we tested for how such size biases in turn translate into differences among populations in the sizes of the fish. Bears caught larger-than-average salmon as the spawning season progressed and as predicted, this was most pronounced in streams with fewer refugia for the fish (i.e., wood and undercut banks). Salmon were marginally smaller in streams with more pronounced size-biased predation but this predictor was less reliable than physical characteristics of streams, with larger fish in wider, deeper streams. These results support the hypothesis that selective forces imposed by predators can be mediated by habitat characteristics, with potential consequences for physical traits of prey.