Metabolic constraints and individual variation shape the trade-off between physiological recovery and anti-predator responses in adult sockeye salmon.

Metabolic scope represents the aerobic energy budget available to an organism to perform non-maintenance activities (e.g., escape a predator, recover from a fisheries interaction, compete for a mate). Conflicting energetic requirements can give rise to ecologically relevant metabolic trade-offs when energy budgeting is constrained. The objective of this study was to investigate how aerobic energy is utilized when individual sockeye salmon (Oncorhynchus nerka) are exposed to multiple acute stressors. To indirectly assess metabolic changes in free-swimming individuals, salmon were implanted with heart rate biologgers. The animals were then exercised to exhaustion or briefly handled as a control, and allowed to recover from this stressor for 48 h. During the first 2 h of the recovery period, individual salmon were exposed to 90 ml of conspecific alarm cues or water as a control. Heart rate was recorded throughout the recovery period. Recovery effort and time was higher in exercised fish, relative to control fish, whereas exposure to an alarm cue had no effect on either of these metrics. Individual routine heart rate was negatively correlated with recovery time and effort. Together, these findings suggest that metabolic energy allocation towards exercise recovery (i.e., an acute stressor; handling, chase, etc.) trumps anti-predator responses in salmon, although individual variation may mediate this effect at the population level.

Infected juvenile salmon can experience increased predation during freshwater migration.

Predation risk for animal migrants can be impacted by physical condition. Although size- or condition-based selection is often observed, observing infection-based predation is rare due to the difficulties in assessing infectious agents in predated samples. We examined predation of outmigrating sockeye salmon (Oncorhynchus nerka) smolts by bull trout (Salvelinus confluentus) in south-central British Columbia, Canada. We used a high-throughput quantitative polymerase chain reaction (qPCR) platform to screen for the presence of 17 infectious agents found in salmon and assess 14 host genes associated with viral responses. In one (2014) of the two years assessed (2014 and 2015), the presence of infectious haematopoietic necrosis virus (IHNv) resulted in 15-26 times greater chance of predation; in 2015 IHNv was absent among all samples, predated or not. Thus, we provide further evidence that infection can impact predation risk in migrants. Some smolts with high IHNv loads also exhibited gene expression profiles consistent with a virus-induced disease state. Nine other infectious agents were observed between the two years, none of which were associated with increased selection by bull trout. In 2014, richness of infectious agents was also associated with greater predation risk. This is a rare demonstration of predator consumption resulting in selection for prey that carry infectious agents. The mechanism by which this selection occurs is not yet determined. By culling infectious agents from migrant populations, fish predators could provide an ecological benefit to prey.

Language Specificity in Phonetic Cue Weighting: Monolingual and Bilingual Perception of the Stop Voicing Contrast in English and Spanish.

BACKGROUND/AIMS: This work examines the perception of the stop voicing contrast in Spanish and English along four acoustic dimensions, comparing monolingual and bilingual listeners. Our primary goals are to test the extent to which cue-weighting strategies are language-specific in monolinguals, and whether this language specificity extends to bilingual listeners. METHODS: Participants categorized sounds varying in voice onset time (VOT, the primary cue to the contrast) and three secondary cues: fundamental frequency at vowel onset, first formant (F1) onset frequency, and stop closure duration. Listeners heard acoustically identical target stimuli, within language-specific carrier phrases, in English and Spanish modes. RESULTS: While all listener groups used all cues, monolingual English listeners relied more on F1, and less on closure duration, than monolingual Spanish listeners, indicating language specificity in cue use. Early bilingual listeners used the three secondary cues similarly in English and Spanish, despite showing language-specific VOT boundaries. CONCLUSION: While our findings reinforce previous work demonstrating language-specific phonetic representations in bilinguals in terms of VOT boundary, they suggest that this specificity may not extend straightforwardly to cue-weighting strategies.

How bilinguals perceive speech depends on which language they think they're hearing.

Bilinguals understand when the communication context calls for speaking a particular language and can switch from speaking one language to speaking the other based on such conceptual knowledge. There is disagreement regarding whether conceptually-based language selection is also possible in the listening modality. For example, can bilingual listeners perceptually adjust to changes in pronunciation across languages based on their conceptual understanding of which language they're currently hearing? We asked French- and Spanish-English bilinguals to identify nonsense monosyllables as beginning with /b/ or /p/, speech categories that French and Spanish speakers pronounce differently than English speakers. We conceptually cued each bilingual group to one of their two languages or the other by explicitly instructing them that the speech items were word onsets in that language, uttered by a native speaker thereof. Both groups adjusted their /b-p/ identification boundary as a function of this conceptual cue to the language context. These results support a bilingual model permitting conceptually-based language selection on both the speaking and listening end of a communicative exchange.

Procedural-Memory, Working-Memory, and Declarative-Memory Skills Are Each Associated With Dimensional Integration in Sound-Category Learning.

This paper investigates relationships between procedural-memory, declarative-memory, and working-memory skills and adult native English speakers' novel sound-category learning. Participants completed a sound-categorization task that required integrating two dimensions: one native (vowel quality), one non-native (pitch). Similar information-integration category structures in the visual and auditory domains have been shown to be best learned implicitly (e.g., Maddox et al., 2006). Thus, we predicted that individuals with greater procedural-memory capacity would better learn sound categories, because procedural memory appears to support implicit learning of new information and integration of dimensions. Seventy undergraduates were tested across two experiments. Procedural memory was assessed using a linguistic adaptation of the serial-reaction-time task (Misyak et al., 2010a,b). Declarative memory was assessed using the logical-memory subtest of the Wechsler Memory Scale-4th edition (WMS-IV; Wechsler, 2009). Working memory was assessed using an auditory version of the reading-span task (Kane et al., 2004). Experiment 1 revealed contributions of only declarative memory to dimensional integration, which might indicate not enough time or motivation to shift over to a procedural/integrative strategy. Experiment 2 gave twice the speech-sound training, distributed over 2 days, and also attempted to train at the category boundary. As predicted, effects of declarative memory were removed and effects of procedural memory emerged, but, unexpectedly, new effects of working memory surfaced. The results may be compatible with a multiple-systems account in which declarative and working memory facilitate transfer of control to the procedural system.

Tracking wild sockeye salmon smolts to the ocean reveals distinct regions of nocturnal movement and high mortality.

Few estimates of migration rates or descriptions of behavior or survival exist for wild populations of out-migrating Pacific salmon smolts from natal freshwater rearing areas to the ocean. Using acoustic transmitters and fixed receiver arrays across four years (2010-2013), we tracked the migration of > 1850 wild sockeye salmon (Oncorhynchus nerka) smolts from Chilko Lake, British Columbia, to the coastal Pacific Ocean (> 1000 km distance). Cumulative survival to the ocean ranged 3-10% among years, although this may be slightly underestimated due to technical limitations at the final receiver array. Distinct spatial patterns in both behavior and survival were observed through all years. In small, clear, upper-river reaches, downstream migration largely occurred at night at speeds up to 50 km/d and coincided with poor survival. Among years, only 57-78% of smolts survived the first 80 km. Parallel laboratory experiments revealed excellent short-term survival and unhindered swimming performance of dummy-tagged smolts, suggesting that predators rather than tagging effects were responsible for the initial high mortality of acoustic-tagged smolts. Migration speeds increased in the Fraser River mainstem (~220 km/d in some years), diel movement patterns ceased, and smolt survival generally exceeded 90% in this segment. Marine movement rates and survival were variable across years, with among-year segment-specific survival being the most variable and lowest (19-61%) during the final (and longest, 240 km) marine migration segment. Osmoregulatory preparedness was not expected to influence marine survival, as smolts could maintain normal levels of plasma chloride when experimentally exposed to saltwater (30 ppt) immediately upon commencing their migration from Chilko Lake. Transportation of smolts downstream generally increased survival to the farthest marine array. The act of tagging may have affected smolts in the marine environment in some years as dummy-tagged fish had poorer survival than control fish when transitioned to saltwater in laboratory-based experiments. Current fisheries models for forecasting the number of adult sockeye returning to spawn have been inaccurate in recent years and generally do not incorporate juvenile or smolt survival information. Our results highlight significant potential for early migration conditions to influence adult recruitment.

Predator swamping reduces predation risk during nocturnal migration of juvenile salmon in a high-mortality landscape.

Animal migrations are costly and are often characterized by high predation risk for individuals. Three of the most oft-assumed mechanisms for reducing risk for migrants are swamping predators with high densities, specific timing of migrations and increased body size. Assessing the relative importance of these mechanisms in reducing predation risk particularly for migrants is generally lacking due to the difficulties in tracking the fate of individuals and population-level characteristics simultaneously. We used acoustic telemetry to track migration behaviour and survival of juvenile sockeye salmon (Oncorhynchus nerka) smolts released over a wide range of conspecific outmigration densities in a river associated with poor survival. The landscape was indeed high risk; smolt survival was poor (˜68%) over 13·5 km of river examined even though migration was rapid (generally <48 h). Our results demonstrate that smolts largely employ swamping of predators to reduce predation risk. Increased densities of co-migrant conspecifics dramatically improved survival of smolts. The strong propensity for nocturnal migration resulted in smolts pausing downstream movements until the next nightfall, greatly increasing relative migration durations for smolts that could not traverse the study area in a single night. Smolt size did not appear to impact predation risk, potentially due to unique characteristics of the system or our inability to tag the entire size range of outmigrants. Movement behaviours were important in traversing this high-risk landscape and provide rare evidence for swamping to effectively reduce individual predation risk.

Individual differences in perceptual adaptability of foreign sound categories.

Listeners possess a remarkable ability to adapt to acoustic variability in the realization of speech sound categories (e.g., different accents). The current work tests whether non-native listeners adapt their use of acoustic cues in phonetic categorization when they are confronted with changes in the distribution of cues in the input, as native listeners do, and examines to what extent these adaptation patterns are influenced by individual cue-weighting strategies. In line with previous work, native English listeners, who use voice onset time (VOT) as a primary cue to the stop voicing contrast (e.g., 'pa' vs. 'ba'), adjusted their use of f0 (a secondary cue to the contrast) when confronted with a noncanonical "accent" in which the two cues gave conflicting information about category membership. Native Korean listeners' adaptation strategies, while variable, were predictable based on their initial cue weighting strategies. In particular, listeners who used f0 as the primary cue to category membership adjusted their use of VOT (their secondary cue) in response to the noncanonical accent, mirroring the native pattern of "downweighting" a secondary cue. Results suggest that non-native listeners show native-like sensitivity to distributional information in the input and use this information to adjust categorization, just as native listeners do, with the specific trajectory of category adaptation governed by initial cue-weighting strategies.

Individual differences in phonetic cue use in production and perception of a non-native sound contrast.

The current work examines native Korean speakers' perception and production of stop contrasts in their native language (L1, Korean) and second language (L2, English), focusing on three acoustic dimensions that are all used, albeit to different extents, in both languages: voice onset time (VOT), f0 at vowel onset, and closure duration. Participants used all three cues to distinguish the L1 Korean three-way stop distinction in both production and perception. Speakers' productions of the L2 English contrasts were reliably distinguished using both VOT and f0 (even though f0 is only a very weak cue to the English contrast), and, to a lesser extent, closure duration. In contrast to the relative homogeneity of the L2 productions, group patterns on a forced-choice perception task were less clear-cut, due to considerable individual differences in perceptual categorization strategies, with listeners using either primarily VOT duration, primarily f0, or both dimensions equally to distinguish the L2 English contrast. Differences in perception, which were stable across experimental sessions, were not predicted by individual variation in production patterns. This work suggests that reliance on multiple cues in representation of a phonetic contrast can form the basis for distinct individual cue-weighting strategies in phonetic categorization.

Phoneme categorization relying solely on high-frequency energy.

Speech perception studies generally focus on the acoustic information present in the frequency regions below 6 kHz. Recent evidence suggests that there is perceptually relevant information in the higher frequencies, including information affecting speech intelligibility. This experiment examined whether listeners are able to accurately identify a subset of vowels and consonants in CV-context when only high-frequency (above 5 kHz) acoustic information is available (through high-pass filtering and masking of lower frequency energy). The findings reveal that listeners are capable of extracting information from these higher frequency regions to accurately identify certain consonants and vowels.

Discriminating simulated vocal tremor source using amplitude modulation spectra.

OBJECTIVES/HYPOTHESIS: Sources of vocal tremor are difficult to categorize perceptually and acoustically. This article describes a preliminary attempt to discriminate vocal tremor sources through the use of spectral measures of the amplitude envelope. The hypothesis is that different vocal tremor sources are associated with distinct patterns of acoustic amplitude modulations. STUDY DESIGN: Statistical categorization methods (discriminant function analysis) were used to discriminate signals from simulated vocal tremor with different sources using only acoustic measures derived from the amplitude envelopes. METHODS: Simulations of vocal tremor were created by modulating parameters of a vocal fold model corresponding to oscillations of respiratory driving pressure (respiratory tremor), degree of vocal fold adduction (adductory tremor), and fundamental frequency of vocal fold vibration (F0 tremor). The acoustic measures were based on spectral analyses of the amplitude envelope computed across the entire signal and within select frequency bands. RESULTS: The signals could be categorized (with accuracy well above chance) in terms of the simulated tremor source using only measures of the amplitude envelope spectrum even when multiple sources of tremor were included. CONCLUSIONS: These results supply initial support for an amplitude-envelope-based approach to identify the source of vocal tremor and provide further evidence for the rich information about talker characteristics present in the temporal structure of the amplitude envelope.

Gender and vocal production mode discrimination using the high frequencies for speech and singing.

Humans routinely produce acoustical energy at frequencies above 6 kHz during vocalization, but this frequency range is often not represented in communication devices and speech perception research. Recent advancements toward high-definition (HD) voice and extended bandwidth hearing aids have increased the interest in the high frequencies. The potential perceptual information provided by high-frequency energy (HFE) is not well characterized. We found that humans can accomplish tasks of gender discrimination and vocal production mode discrimination (speech vs. singing) when presented with acoustic stimuli containing only HFE at both amplified and normal levels. Performance in these tasks was robust in the presence of low-frequency masking noise. No substantial learning effect was observed. Listeners also were able to identify the sung and spoken text (excerpts from "The Star-Spangled Banner") with very few exposures. These results add to the increasing evidence that the high frequencies provide at least redundant information about the vocal signal, suggesting that its representation in communication devices (e.g., cell phones, hearing aids, and cochlear implants) and speech/voice synthesizers could improve these devices and benefit normal-hearing and hearing-impaired listeners.

Immune response genes and pathogen presence predict migration survival in wild salmon smolts.

We present the first data to link physiological responses and pathogen presence with subsequent fate during migration of wild salmonid smolts. We tagged and non-lethally sampled gill tissue from sockeye salmon (Oncorhynchus nerka) smolts as they left their nursery lake (Chilko Lake, BC, Canada) to compare gene expression profiles and freshwater pathogen loads with migration success over the first ~1150 km of their migration to the North Pacific Ocean using acoustic telemetry. Fifteen per cent of smolts were never detected again after release, and these fish had gene expression profiles consistent with an immune response to one or more viral pathogens compared with fish that survived their freshwater migration. Among the significantly upregulated genes of the fish that were never detected postrelease were MX (interferon-induced GTP-binding protein Mx) and STAT1 (signal transducer and activator of transcription 1-alpha/beta), which are characteristic of a type I interferon response to viral pathogens. The most commonly detected pathogen in the smolts leaving the nursery lake was infectious haematopoietic necrosis virus (IHNV). Collectively, these data show that some of the fish assumed to have died after leaving the nursery lake appeared to be responding to one or more viral pathogens and had elevated stress levels that could have contributed to some of the mortality shortly after release. We present the first evidence that changes in gene expression may be predictive of some of the freshwater migration mortality in wild salmonid smolts.

The perceptual significance of high-frequency energy in the human voice.

While human vocalizations generate acoustical energy at frequencies up to (and beyond) 20 kHz, the energy at frequencies above about 5 kHz has traditionally been neglected in speech perception research. The intent of this paper is to review (1) the historical reasons for this research trend and (2) the work that continues to elucidate the perceptual significance of high-frequency energy (HFE) in speech and singing. The historical and physical factors reveal that, while HFE was believed to be unnecessary and/or impractical for applications of interest, it was never shown to be perceptually insignificant. Rather, the main causes for focus on low-frequency energy appear to be because the low-frequency portion of the speech spectrum was seen to be sufficient (from a perceptual standpoint), or the difficulty of HFE research was too great to be justifiable (from a technological standpoint). The advancement of technology continues to overcome concerns stemming from the latter reason. Likewise, advances in our understanding of the perceptual effects of HFE now cast doubt on the first cause. Emerging evidence indicates that HFE plays a more significant role than previously believed, and should thus be considered in speech and voice perception research, especially in research involving children and the hearing impaired.

Acoustic and perceptual effects of left-right laryngeal asymmetries based on computational modeling.

PURPOSE: Computational modeling was used to examine the consequences of 5 different laryngeal asymmetries on acoustic and perceptual measures of vocal function. METHOD: A kinematic vocal fold model was used to impose 5 laryngeal asymmetries: adduction, edge bulging, nodal point ratio, amplitude of vibration, and starting phase. Thirty /a/ and /ɪ/ vowels were generated for each asymmetry and analyzed acoustically using cepstral peak prominence (CPP), harmonics-to-noise ratio (HNR), and 3 measures of spectral slope (H1*-H2*, B0-B1, and B0-B2). Twenty listeners rated voice quality for a subset of the productions. RESULTS: Increasingly asymmetric adduction, bulging, and nodal point ratio explained significant variance in perceptual rating (R2 = .05, p < .001). The same factors resulted in generally decreasing CPP, HNR, and B0-B2 and in increasing B0-B1. Of the acoustic measures, only CPP explained significant variance in perceived quality (R2 = .14, p < .001). Increasingly asymmetric amplitude of vibration or starting phase minimally altered vocal function or voice quality. CONCLUSION: Asymmetries of adduction, bulging, and nodal point ratio drove acoustic measures and perception in the current study, whereas asymmetric amplitude of vibration and starting phase demonstrated minimal influence on the acoustic signal or voice quality.

The alluring but misleading analogy between mirror neurons and the motor theory of speech.

Speech is commonly claimed to relate to mirror neurons because of the alluring surface analogy of mirror neurons to the Motor Theory of speech perception, which posits that perception and production draw upon common motor-articulatory representations. We argue that the analogy fails and highlight examples of systems-level developmental approaches that have been more fruitful in revealing perception-production associations.

Disentangling the roles of air exposure, gill net injury, and facilitated recovery on the postcapture and release mortality and behavior of adult migratory sockeye salmon (Oncorhynchus nerka) in freshwater.

We sought to improve the understanding of delayed mortality in migrating sockeye salmon (Oncorhynchus nerka) captured and released in freshwater fisheries. Using biotelemetry, blood physiology, and reflex assessments, we evaluated the relative roles of gill net injury and air exposure and investigated whether using a recovery box improved survival. Fish (n=238), captured by beach seine, were allocated to four treatment groups: captured only, air exposed, injured, and injured and air exposed. Only half of the fish in each group were provided with a 15-min facilitated recovery. After treatment, fish were radio-tagged and released to resume their migration. Blood status was assessed in 36 additional untagged fish sampled after the four treatments. Compared with fish sampled immediately on capture, all treatments resulted in elevated plasma lactate and cortisol concentrations. After air exposure, plasma osmolality was elevated and reflexes were significantly impaired relative to the control and injured treatments. Injured fish exhibited reduced short-term migration speed by 3.2 km/d and had a 14.5% reduced survival to subnatal watersheds compared to controls. The 15-min facilitated recovery improved reflex assessment relative to fish released immediately but did not affect survival. We suggest that in sockeye salmon migrating in cool water temperatures (∼13°-16°C), delayed mortality can result from injury and air exposure, perhaps through sublethal stress, and that injury created additive delayed mortality likely via secondary infections.

Detection of high-frequency energy level changes in speech and singing.

Previous work has shown that human listeners are sensitive to level differences in high-frequency energy (HFE) in isolated vowel sounds produced by male singers. Results indicated that sensitivity to HFE level changes increased with overall HFE level, suggesting that listeners would be more "tuned" to HFE in vocal production exhibiting higher levels of HFE. It follows that sensitivity to HFE level changes should be higher (1) for female vocal production than for male vocal production and (2) for singing than for speech. To test this hypothesis, difference limens for HFE level changes in male and female speech and singing were obtained. Listeners showed significantly greater ability to detect level changes in singing vs speech but not in female vs male speech. Mean differences limen scores for speech and singing were about 5 dB in the 8-kHz octave (5.6-11.3 kHz) but 8-10 dB in the 16-kHz octave (11.3-22 kHz). These scores are lower (better) than those previously reported for isolated vowels and some musical instruments.

A Bafri, un Pafri: bilinguals' Pseudoword identifications support language-specific phonetic systems.

Bilinguals perceptually accommodate speech variation across languages, but to what extent this flexibility depends on bilingual experience is uncertain. One account suggests that bilingual experience promotes language-specific processing modes, implying that bilinguals can switch as appropriate between the different phonetic systems of the languages they speak. Another account suggests that bilinguals rapidly recalibrate to the unique acoustic properties of each language following language-general processes common to monolinguals. Challenging this latter account, the present results show that Spanish-English bilinguals with exposure to both languages from early childhood, but not English monolinguals, shift perception as appropriate across acoustically controlled English and Spanish contexts. Early bilingual experience appears to promote language-specific phonetic systems.