Assortative mate preferences for height across short-term and long-term relationship contexts in a cross-cultural sample.

Height preferences reflecting positive assortative mating for height-wherein an individual's own height positively predicts the preferred height of their mate-have been observed in several distinct human populations and are thought to increase reproductive fitness. However, the extent to which assortative preferences for height differ strategically for short-term versus long-term relationship partners, as they do for numerous other indices of mate quality, remains unclear. We explore this possibility in a large representative sample of over 500 men and women aged 15-77 from Canada, Cuba, Norway and the United States. Participants' own heights were measured, and they indicated their height preferences for a long-term and short-term mate using graphic stimuli containing metric indices. Replicating the "male-taller norm," participants on average preferred taller-than-average male mates, and shorter-than-average female mates. Positive assortative preferences for height were observed across sexes and samples, however the strength of these height preferences varied with relationship context for men, and not for women. Taller men preferred relatively shorter women for short-term relationships than for long-term relationships, indicating stronger assortative preferences for height in a long-term context. These results provide preliminary evidence that, in addition to mate preferences for other physical traits related to mate quality such as masculinity in the body, face, and voice, assortative preferences for height do vary as a function of expected relationship length, but this was surprisingly only observed in preferences for female height.

Hearing diversity in moths confronting a neotropical bat assemblage.

The tympanal ear is an evolutionary acquisition which helps moths survive predation from bats. The greater diversity of bats and echolocation strategies in the Neotropics compared with temperate zones would be expected to impose different sensory requirements on the neotropical moths. However, even given some variability among moth assemblages, the frequencies of best hearing of moths from different climate zones studied to date have been roughly the same: between 20 and 60 kHz. We have analyzed the auditory characteristics of tympanate moths from Cuba, a neotropical island with high levels of bat diversity and a high incidence of echolocation frequencies above those commonly at the upper limit of moths' hearing sensitivity. Moths of the superfamilies Noctuoidea, Geometroidea and Pyraloidea were examined. Audiograms were determined by non-invasively measuring distortion-product otoacoustic emissions. We also quantified the frequency spectrum of the echolocation sounds to which this moth community is exposed. The hearing ranges of moths in our study showed best frequencies between 36 and 94 kHz. High sensitivity to frequencies above 50 kHz suggests that the auditory sensitivity of moths is suited to the sounds used by sympatric echolocating bat fauna. Biodiversity characterizes predators and prey in the Neotropics, but the bat-moth acoustic interaction keeps spectrally matched.

Volitional exaggeration of body size through fundamental and formant frequency modulation in humans.

Several mammalian species scale their voice fundamental frequency (F0) and formant frequencies in competitive and mating contexts, reducing vocal tract and laryngeal allometry thereby exaggerating apparent body size. Although humans' rare capacity to volitionally modulate these same frequencies is thought to subserve articulated speech, the potential function of voice frequency modulation in human nonverbal communication remains largely unexplored. Here, the voices of 167 men and women from Canada, Cuba, and Poland were recorded in a baseline condition and while volitionally imitating a physically small and large body size. Modulation of F0, formant spacing (∆F), and apparent vocal tract length (VTL) were measured using Praat. Our results indicate that men and women spontaneously and systemically increased VTL and decreased F0 to imitate a large body size, and reduced VTL and increased F0 to imitate small size. These voice modulations did not differ substantially across cultures, indicating potentially universal sound-size correspondences or anatomical and biomechanical constraints on voice modulation. In each culture, men generally modulated their voices (particularly formants) more than did women. This latter finding could help to explain sexual dimorphism in F0 and formants that is currently unaccounted for by sexual dimorphism in human vocal anatomy and body size.

Echo-level compensation and delay tuning in the auditory cortex of the mustached bat.

During echolocation, bats continuously perform audio-motor adjustments to optimize detection efficiency. It has been demonstrated that bats adjust the amplitude of their biosonar vocalizations (known as 'pulses') to stabilize the amplitude of the returning echo. Here, we investigated this echo-level compensation behaviour by swinging mustached bats on a pendulum towards a reflective surface. In such a situation, the bats lower the amplitude of their emitted pulses to maintain the amplitude of incoming echoes at a constant level as they approach a target. We report that cortical auditory neurons that encode target distance have receptive fields that are optimized for dealing with echo-level compensation. In most cortical delay-tuned neurons, the echo amplitude eliciting the maximum response matches the echo amplitudes measured from the bats' biosonar vocalizations while they are swung in a pendulum. In addition, neurons tuned to short target distances are maximally responsive to low pulse amplitudes while neurons tuned to long target distances respond maximally to high pulse amplitudes. Our results suggest that bats dynamically adjust biosonar pulse amplitude to match the encoding of target range and to keep the amplitude of the returning echo within the bounds of the cortical map of echo delays.

Unexpected dynamic up-tuning of auditory organs in day-flying moths.

In certain nocturnal moth species the frequency range of best hearing shifts to higher frequencies during repeated sound stimulation. This could provide the moths with a mechanism to better detect approaching echolocating bats. However, such a dynamic up-tuning would be of little value for day-flying moths that use intra-specific acoustic communication. Here we examined if the ears of day-flying moths provide stable tuning during longer sound stimulation. Contrary to our expectations, dynamic up-tuning was found in the ear of the day-flying species Urania boisduvalii and Empyreuma pugione. Audiograms were measured with distortion-product otoacoustic emissions (DPOAEs). The level of the dominant distortion product (i.e. 2f1-f2) varied as a function of time by as much as 45 dB during ongoing acoustic stimulation, showing a systematic decrease at low frequencies and an increase at high frequencies. As a consequence, within about 2 s of acoustic stimulation, the DPOAEs audiogram shifted from low to high frequencies. Despite the up-tuning, the range of best audition still fell within the frequency band of the species-specific communication signals, suggesting that intra-specific communication should not be affected adversely. Up-tuning could be an ancestral condition in moth ears that in day-flying moths does not underlie larger selection pressure.

Level-tolerant duration selectivity in the auditory cortex of the velvety free-tailed bat Molossus molossus.

It has been reported previously that in the inferior colliculus of the bat Molossus molossus, neuronal duration tuning is ambiguous because the tuning type of the neurons dramatically changes with the sound level. In the present study, duration tuning was examined in the auditory cortex of M. molossus to describe if it is as ambiguous as the collicular tuning. From a population of 174 cortical 104 (60 %) neurons did not show duration selectivity (all-pass). Around 5 % (9 units) responded preferentially to stimuli having longer durations showing long-pass duration response functions, 35 (20 %) responded to a narrow range of stimulus durations showing band-pass duration response functions, 24 (14 %) responded most strongly to short stimulus durations showing short-pass duration response functions and two neurons (1 %) responded best to two different stimulus durations showing a two-peaked duration-response function. The majority of neurons showing short- (16 out of 24) and band-pass (24 out 35) selectivity displayed "O-shaped" duration response areas. In contrast to the inferior colliculus, duration tuning in the auditory cortex of M. molossus appears level tolerant. That is, the type of duration selectivity and the stimulus duration eliciting the maximum response were unaffected by changing sound level.

Tone-deaf ears in moths may limit the acoustic detection of two-tone bats.

Frequency alternation in the echolocation of insectivorous bats has been interpreted in relation to ranging and duty cycle, i.e. advantages for echolocation. The shifts in frequency of the calls of these so-called two-tone bats, however, may also play its role in the success of their hunting behavior for a preferred prey, the tympanate moth. How the auditory receptors (e.g. the A1 and A2 cells) in the moth's ear detect such frequency shifts is currently unknown. Here, we measured the auditory responses of the A1 cell in the noctuid Spodoptera frugiperda to the echolocation hunting sequence of Molossus molossus, a two-tone bat. We also manipulated the bat calls to control for the frequency shifts by lowering the frequency band of the search and approach calls. The firing response of the A1 receptor cell significantly decreases with the shift to higher frequencies during the search and approach phases of the hunting sequence of M. molossus; this could be explained by the receptor's threshold curve. The frequency dependence of the decrease in the receptor's response is supported by the results attained with the manipulated sequence: search and approach calls with the same minimum frequency are detected by the moth at the same threshold intensity. The two-tone bat M. molossus shows a call frequency alternation behavior that may enable it to overcome moth audition even in the mid-frequency range (i.e. 20-50 kHz) where moths hear best.

Narrow sound pressure level tuning in the auditory cortex of the bats Molossus molossus and Macrotus waterhousii.

In the auditory system, tuning to sound level appears in the form of non-monotonic response-level functions that depict the response of a neuron to changing sound levels. Neurons with non-monotonic response-level functions respond best to a particular sound pressure level (defined as "best level" or level evoking the maximum response). We performed a comparative study on the location and basic functional organization of the auditory cortex in the gleaning bat, Macrotus waterhousii, and the aerial-hawking bat, Molossus molossus. Here, we describe the response-level function of cortical units in these two species. In the auditory cortices of M. waterhousii and M. molossus, the characteristic frequency of the units increased from caudal to rostral. In M. waterhousii, there was an even distribution of characteristic frequencies while in M. molossus there was an overrepresentation of frequencies present within echolocation pulses. In both species, most of the units showed best levels in a narrow range, without an evident topography in the amplitopic organization, as described in other species. During flight, bats decrease the intensity of their emitted pulses when they approach a prey item or an obstacle resulting in maintenance of perceived echo intensity. Narrow level tuning likely contributes to the extraction of echo amplitudes facilitating echo-intensity compensation. For aerial-hawking bats, like M. molossus, receiving echoes within the optimal sensitivity range can help the bats to sustain consistent analysis of successive echoes without distortions of perception caused by changes in amplitude.

Blurry topography for precise target-distance computations in the auditory cortex of echolocating bats.

Echolocating bats use the time from biosonar pulse emission to the arrival of echo (defined as echo delay) to calculate the space depth of targets. In the dorsal auditory cortex of several species, neurons that encode increasing echo delays are organized rostrocaudally in a topographic arrangement defined as chronotopy. Precise chronotopy could be important for precise target-distance computations. Here we show that in the cortex of three echolocating bat species (Pteronotus quadridens, Pteronotus parnellii and Carollia perspicillata), chronotopy is not precise but blurry. In all three species, neurons throughout the chronotopic map are driven by short echo delays that indicate the presence of close targets and the robustness of map organization depends on the parameter of the receptive field used to characterize neuronal tuning. The timing of cortical responses (latency and duration) provides a binding code that could be important for assembling acoustic scenes using echo delay information from objects with different space depths.

Mechanical tuning of the moth ear: distortion-product otoacoustic emissions and tympanal vibrations.

The mechanical tuning of the ear in the moth Empyreuma pugione was investigated by distortion-product otoacoustic emissions (DPOAE) and laser Doppler vibrometry (LDV). DPOAE audiograms were assessed using a novel protocol that may be advantageous for non-invasive auditory studies in insects. To evoke DPOAE, two-tone stimuli within frequency and level ranges that generated a large matrix of values (960 frequency-level combinations) were used to examine the acoustic space in which the moth tympanum shows its best mechanical and acoustical responses. The DPOAE tuning curve derived from the response matrix resembles that obtained previously by electrophysiology, and is V-shaped and tuned to frequencies between 25 and 45 kHz with low Q10dB values of 1.21±0.26. In addition, while using a comparable stimulation regime, mechanical distortion in the displacement of the moth's tympanal membrane at the stigma was recorded with a laser Doppler vibrometer. The corresponding mechanical vibration audiograms were compared with DPOAE audiograms. Both types of audiograms have comparable shape, but most of the mechanical response fields are shifted towards lower frequencies. We showed for the first time in moths that DPOAE have a pronounced analogy in the vibration of the tympanic membrane where they may originate. Our work supports previous studies that point to the stigma (and the internally associated transduction machinery) as an important place of sound amplification in the moth ear, but also suggests a complex mechanical role for the rest of the transparent zone.

Evolution of the heteroharmonic strategy for target-range computation in the echolocation of Mormoopidae.

Echolocating bats use the time elapsed from biosonar pulse emission to the arrival of echo (defined as echo-delay) to assess target-distance. Target-distance is represented in the brain by delay-tuned neurons that are classified as either "heteroharmonic" or "homoharmormic." Heteroharmonic neurons respond more strongly to pulse-echo pairs in which the timing of the pulse is given by the fundamental biosonar harmonic while the timing of echoes is provided by one (or several) of the higher order harmonics. On the other hand, homoharmonic neurons are tuned to the echo delay between similar harmonics in the emitted pulse and echo. It is generally accepted that heteroharmonic computations are advantageous over homoharmonic computations; i.e., heteroharmonic neurons receive information from call and echo in different frequency-bands which helps to avoid jamming between pulse and echo signals. Heteroharmonic neurons have been found in two species of the family Mormoopidae (Pteronotus parnellii and Pteronotus quadridens) and in Rhinolophus rouxi. Recently, it was proposed that heteroharmonic target-range computations are a primitive feature of the genus Pteronotus that was preserved in the evolution of the genus. Here, we review recent findings on the evolution of echolocation in Mormoopidae, and try to link those findings to the evolution of the heteroharmonic computation strategy (HtHCS). We stress the hypothesis that the ability to perform heteroharmonic computations evolved separately from the ability of using long constant-frequency echolocation calls, high duty cycle echolocation, and Doppler Shift Compensation. Also, we present the idea that heteroharmonic computations might have been of advantage for categorizing prey size, hunting eared insects, and living in large conspecific colonies. We make five testable predictions that might help future investigations to clarify the evolution of the heteroharmonic echolocation in Mormoopidae and other families.

Neurons in the inferior colliculus of the mustached bat are tuned both to echo-delay and sound duration.

Echolocation in bats requires a precise temporal processing of complex signals. This processing of time includes the encoding of echo-delay, which gives an estimation of target distance, and sound duration, which is considered to be important for own sound or echo recognition. In this study, we report that delay-tuned neurons in the inferior colliculus of the mustached bat (Pteronotus parnellii) are also tuned to sound duration. Collicular delay-tuned neurons showed three types of duration tuning: short-pass (12 of 64), band-pass (16 of 64), and long-pass (17 of 64). The remaining 19 delay-tuned neurons are not selective for sound duration. All short-pass and 10 band-pass neurons' characteristic delays were similar to characteristic duration. In six band-pass neurons, characteristic delay was different from characteristic duration. Neurons processing unmatched delay and durations could be participating in complex kinds of processing where the same neuron has different functions depending on the activated neural network.

Evolution of neuronal mechanisms for echolocation: specializations for target-range computation in bats of the genus Pteronotus.

Delay tuning was studied in the auditory cortex of Pteronotus quadridens. All the 136 delay-tuned units that were studied responded strongly to heteroharmonic pulse-echo pairs presented at specific delays. In the heteroharmonic pairs, the first sonar call harmonic marks the timing of pulse emission while one of the higher harmonics (second or third) indicates the timing of the echo. Delay-tuned units are organized chronotopically along a rostrocaudal axis according to their characteristic delay. There is no obvious indication of multiple cortical axes specialized in the processing of different harmonic combinations of pulse and echo. Results of this study serve for a straight comparison of cortical delay-tuning between P. quadridens and the well-studied mustached bat, Pteronotus parnellii. These two species stem from the most recent and most basal nodes in the Pteronotus lineage, respectively. P. quadridens and P. parnellii use comparable heteroharmonic target-range computation strategies even though they do not use biosonar calls of a similar design. P. quadridens uses short constant-frequency (CF)/frequency-modulated (FM) echolocation calls, while P. parnellii uses long CF/FM calls. The ability to perform "heteroharmonic" target-range computations might be an ancestral neuronal specialization of the genus Pteronotus that was subjected to positive Darwinian selection in the evolution.

Auditory cortex of newborn bats is prewired for echolocation.

Neuronal computation of object distance from echo delay is an essential task that echolocating bats must master for spatial orientation and the capture of prey. In the dorsal auditory cortex of bats, neurons specifically respond to combinations of short frequency-modulated components of emitted call and delayed echo. These delay-tuned neurons are thought to serve in target range calculation. It is unknown whether neuronal correlates of active space perception are established by experience-dependent plasticity or by innate mechanisms. Here we demonstrate that in the first postnatal week, before onset of echolocation and flight, dorsal auditory cortex already contains functional circuits that calculate distance from the temporal separation of a simulated pulse and echo. This innate cortical implementation of a purely computational processing mechanism for sonar ranging should enhance survival of juvenile bats when they first engage in active echolocation behaviour and flight.

Properties of echo delay-tuning receptive fields in the inferior colliculus of the mustached bat.

One role of the inferior colliculus (IC) in bats is to create neuronal delay-tuning, which is used for the estimation of target distance in the echolocating bat's auditory system. In this study, we describe response properties of IC delay-tuned neurons of the mustached bat (Pteronotus parnellii) and compare it with those of delay-tuned neurons of the auditory cortex (AC). We also address the question if frequency content of the stimulus (pure-tone (PT) or frequency-modulated (FM) pairs stimulation) affects combination-sensitive interaction in the same neuron. Sharpness and sensitivity of delay-tuned neurons in the IC are similar to those described in the AC. However, in contrast to cortical responses, in collicular neurons the delay at which the neurons show the maximum response does not change with changes in echo level. This tolerance to changes in the echo level seems to be a property of collicular delay-tuned neurons, which is modified along the ascending auditory pathway. In the IC we found neurons that showed a facilitated delay-tuned response when stimulated with FM components and did not show any delay-tuning with PT stimulation. This result suggests that not only is echo delay-tuning generated in the IC but also its FM-specificity observed in the cortex could be created to some extent in the IC and then topographically organized at higher levels.

Sound-evoked oscillation and paradoxical latency shift in the inferior colliculus neurons of the big fruit-eating bat, Artibeus jamaicensis.

Frequency tuning, temporal response pattern and latency properties of inferior colliculus neurons were investigated in the big fruit-eating bat, Artibeus jamaicensis. Neurons having best frequencies between 48-72 kHz and between 24-32 kHz are overrepresented. The inferior colliculus neurons had either phasic (consisting in only one response cycle at all stimulus intensities) or long-lasting oscillatory responses (consisting of multiple response cycles). Seventeen percent of neurons displayed paradoxical latency shift, i.e. their response latency increased with increasing sound level. Three types of paradoxical latency shift were found: (1) stable, that does not depend on sound duration, (2) duration-dependent, that grows with increasing sound duration, and (3) progressive, whose magnitude increases with increasing sound level. The temporal properties of paradoxical latency shift neurons compare well with those of neurons having long-lasting oscillatory responses, i.e. median inter-spike intervals and paradoxical latency shift below 6 ms are overrepresented. In addition, oscillatory and paradoxical latency shift neurons behave similarly when tested with tones of different durations. Temporal properties of oscillation and PLS found in the IC of fruit-eating bats are similar to those found in the IC of insectivorous bats using downward frequency-modulated echolocation calls.

Duration tuning in the inferior colliculus of the mustached bat.

We studied duration tuning in neurons of the inferior colliculus (IC) of the mustached bat. Duration-tuned neurons in the IC of the mustached bat fall into three main types: short (16 of 136), band (34 of 136), and long (29 of 136) pass. The remaining 51 neurons showed no selectivity for the duration of sounds. The distribution of best durations was double peaked with maxima around 3 and 17 ms, which correlate with the duration of the short frequency-modulated (FM) and the long constant-frequency (CF) signals emitted by Pteronotus parnellii. Since there are no individual neurons with a double-peaked duration response profile, both types of temporal processing seem to be well segregated in the IC. Most short- and band-pass units with best frequency in the CF2 range responded to best durations > 9 ms (66%, 18 of 27 units). However, there is no evidence for a bias toward longer durations as there is for neurons tuned to the frequency range of the FM component of the third harmonic, where 83% (10 of 12 neurons) showed best durations longer than 9 ms. In most duration-tuned neurons, response areas as a function of stimulus duration and intensity showed either V or U shape, with duration tuning retained across the range of sound levels tested. Duration tuning was affected by changes in sound pressure level in only six neurons. In all duration-tuned neurons, latencies measured at the best duration were longer than best durations, suggesting that behavioral decisions based on analysis of the duration of the pulses would not be expected to be complete until well after the stimulus has occurred.

The auditory cortex of the bat Molossus molossus: disproportionate search call frequency representation.

The extent of the auditory cortex in the bat Molossus molossus was electrophysiologically investigated. Best frequencies and minimum thresholds of neural tuning curves were analyzed to define the topography of the auditory cortex. The auditory cortex encompasses an average cortical surface area of 5mm(2). Characteristic frequencies are tonotopically organized with low frequencies being represented caudally and high frequencies rostrally. However, a large interindividual variability in the tonotopic organization was found. In most animals, the caudal 50% was tonotopically organized. More anterior, a variable area was found. A distinct field with reversed topography was not consistently found. Within the demarcated auditory cortex, frequencies of 30-40 kHz, which correspond to the frequency range of search calls emitted during hunting, are overrepresented, occupying 49% of the auditory cortex surface. High minimum thresholds >50 dB SPL were found in a narrow dorsal narrow area. Neurons with multipeaked tuning curves (20%) preferentially were located in the dorsal part of the auditory cortex. In accordance with studies in other bat species, the auditory cortex of M. molossus is highly sensitive to the dominant frequencies of biosonar search calls.

Echolocation in the large molossid bats Eumops glaucinus and Nyctinomops macrotis.

Eumops glaucinus and Nyctinomops macrotis, the largest molossid bats in Cuba, were investigated. Both species of bats share the same guild in the island and are similar in size, which allow the prediction of overlapping echolocation inventories following both the "vocal plasticity hypothesis" and the "scaling hypothesis." In addition, large body size predicts the emission of low frequency calls in the human audible range. Calls recorded during hunting show that the bats' echolocation repertoires are very similar and of low frequency, with most differences in search calls. Matches were found in the calls' design, duration, slope, bandwidth, and spectral parameters. Statistical differences between search calls are consistent with the predictions from the "scaling hypothesis," considering that E. glaucinus is only slightly larger than N. macrotis. The echolocation calls emitted by both species are in the frequency range below 20-25 kHz, which identifies both species as the only ones with echolocation in the human audible range in Cuba.

Echolocation calls of Poey's flower bat (Phyllonycteris poeyi) unlike those of other phyllostomids.

Unlike any other foraging phyllostomid bat studied to date, Poey's flower bats (Phyllonycteris poeyi-Phyllostomidae) emit relatively long (up to 7.2 ms), intense, single-harmonic echolocation calls. These calls are readily detectable at distances of at least 15 m. Furthermore, the echolocation calls contain only the first harmonic, which is usually filtered out in the vocal tract of phyllostomids. The foraging echolocation calls of P. poeyi are more like search-phase echolocation calls of sympatric aerial-feeding bats (Molossidae, Vespertilionidae, Mormoopidae). Intense, long, narrowband, single-harmonic echolocation calls focus acoustic energy maximizing range and favoring detection, which may be particularly important for cruising bats, like P. poeyi, when flying in the open. Flying in enclosed spaces, P. poeyi emit short, low-intensity, frequency-modulated, multiharmonic echolocation calls typical of other phyllostomids. This is the first report of a phyllostomid species emitting long, intense, single-harmonic echolocation calls with most energy in the first harmonic.