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General Psychology

by: Samantha Hettinger

General Psychology PSYC T221

Samantha Hettinger
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This 8 page Class Notes was uploaded by Samantha Hettinger on Saturday September 12, 2015. The Class Notes belongs to PSYC T221 at West Virginia University taught by Staff in Fall. Since its upload, it has received 35 views. For similar materials see /class/202803/psyc-t221-west-virginia-university in Psychlogy at West Virginia University.


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Date Created: 09/12/15
LEVIE 751 Neurobiology of echolocation in bats Cynthia F Moss and Shiva R Sinhai Echolocating bats sub order Microchiroptera form a highly successful group of animals comprising approximately 700 species and an estimated 25 of living mammals Many echolocating bats are nocturnal predators that have evolved a biological sonar system to orient and forage in three dimensional space Acoustic signal processing and vocal motor control are tightly coupled and successful echolocation depends on the coordination between auditory and motor systems Indeed echolocation involves adaptive changes in vocal production patterns which in turn constrain the acoustic information arriving at the bat s ears and the time scales over which neural computations take place Addresses Department of Psychology Institute for Systems Research Neuroscience and Cognitive Science Program University of Maryland College Park MD 20742 USA e mail cmosspsycumdedu e mail sinhawamumdedu Current Opinion in Neurobiology 2003 13751 758 This review comes from a themed issue on Neurobiology of behaviour Edited by Mark Konishi and Randolf Menzel 0959 4388 see front matter 2003 Elsevier Ltd All rights reserved DOI 101016jconb200310016 Abbreviations 3 D three dimensional ACC anterior cingulate cortex BD best duration BF best frequency CF constant frequency DNLL dorsal nucleus of the lateral lemniscus El binaural response profile created with excitatoryinhibitory contralateralipsilateral inputs FM frequency modulated GABA y amino butyric acid IC inferior colliculus ILD interaural level difference PAG periaqueductal gray PB parabrachial nucleus PLa paralemniscal tegmentum area PLS paradoxical latency shift Introduction The echolocating bat s active sensing system supports obstacle avoidance and foraging behavior in complete darkness It produces ultrasonic vocalizations and uses information contained in the returning echoes to deter mine the position size and other features of sonar targets 1 The timing frequency content duration and intensity of sonar signals used by the bat to probe the environment directly in uence the information available to its acoustic imaging system In turn the bat s auditory representation of the environment guides adaptive motor behaviors including adjustments of the pinna head aim ight path and the features of subsequent sonar vocalizations 23 Echolocating bats eXhibit tremendous diversity in the suborder Microchiroptera with species displaying adap tations to a broad range of habitats from the desert to the tropical rain forest 4 Species speci c signal character istics are closely linked to the ecological conditions encountered by foraging bats and several schemes have been proposed to categorize bats according to habitat and sonar signal characteristics 5 9 Each species of bat has a distinct repertoire of signals that it uses for echolocation and the features of these sounds determine the acoustic information available to its sonar imaging system Bat sonar signals fall broadly into two categories constant frequency CF and frequency mod ulated FM see Figure 1a Species using CF FM signals for echolocation typically forage in dense foliage and some of these species adjust the frequency of their sonar vocalizations to compensate for Doppler shifts in return ing echoes 1011 The CF FM bat s Doppler shift compensation DSC serves to cancel a rise in echo frequency introduced by its own ight velocity and iso lates spectral modulations in echoes that come from uttering insect wings 12 In some Doppler shift com pensating bats researchers have identi ed auditory spe cializations which give rise to heightened sensitivity and frequency selectivity in the spectral region of the bat s CF signals 13 By contrast many FM bats forage in the open or at the edge of forests using shorter duration broadband signals that are well suited for three dimen sional 3 D target localization and for separating gure and ground FM bats can discriminate differences in echo delay the cue for target distance of less than 60 micro seconds 114 and they use this delay information to coordinate the timing of sonar vocalizations 15 As an insectiVorous bat ies towards a prey item the spectral temporal features of its sonar vocalizations change Figure 1b The characteristics of sonar emissions have been used to divide the bat s insect pursuit sequence into different phases search approach and terminal buzz 16 These phases of insect capture represent distinct modes of action and perception which provide a valuable system for empirical research on audiomotor feedback control More over the temporal patterning of the bat s echolocation signals provide eXplicit data on the timing of vocal motor wwwcurrent opinioncom Current Opinion in Neurobiology 2003 13751 758 752 Neurobiology of behaviour Figure 1 150 FM M150 Eplesicus fuscus kHz kHz i l l 5 l y l l 0 0 150 CMFM kHz v quotm kqu t l r r 1 HF r v n n 1W5 Approaching prey 50 ms Current Opinion in Neurobiology Echolocation signal structures a Spectrographic examples of frequency modulated FM and constant frequency CF signal components used by echolocating bats b Spectrographic sequence of sI nals produced by an FM bat Eptesicus fuscus and a CF FM bat Rhinoophus ferrumequmum while pursuing insect prey Typical of insectivorous echolocating bats signal repetition rate increases and the duration decreases as the animal approaches its prey commands that feed directly back to the auditory system for spatially guided behavior The echolocating bat s adaptive motor behaviors set a context and timeframe within which neuronal responses must operate and presumably vary to build representa tions of the environment Therefore it is plausible that the timescales over which echolocation behaviors operate serve to constrain the timescales over which neuronal computations take place The potential dynamic variation in neuronal responses can be mediated at the single cell and network level by processes such as experience dependent synaptic plasticity In this review we focus on the temporal parameters of sonar vocalizations and sound processing as they relate to echolocation behavior in bats Auditory processing The bat s auditory system receives and processes echoes in its environment for the task ofspatial orientation but it is essentially a standard mammalian auditory system Many of the same cues used by other species to localize sound and to process complex patterns of acoustic infor mation are exploited by the bat for spatial orientation and perception by sonar Binaural cues for sound localization are used to estimate the azimuthal position of a sonar target The bat s external ear produces changes in the spectrum of incoming echoes which creates patterns of interference that are used by the bat to estimate target elevation 17 The bat estimates the third spatial dimen sion target range from the time delay between the outgoing vocalization and the returning echo 14 and FMbats show extraordinary spatial selectivity along the range axis 1819 Major nuclei comprising the primary auditory pathway and important feedforward and feedback excitatory and inhibitory connections are shown schematically in Figure 2 For a detailed exposition of commonalties and differences in auditory pathways and function in bat species see Fay and Popper 20 Recent research ndings demonstrate that feedback connections can lead to timedependent modi cations in the functional process ing of auditory stimuli 21 This in turn has important consequences for the processing of sequences or combina tions of naturally occurring or arti cially generated stimuli Many neuronal mechanisms operate in the range of 10s to 100s of milliseconds and act in creating response types presumably important in processing echolocation infor mation Mechanisms range from the dependence on the complement of ion currents 22 to the distribution of inhibitory inputs 23 both of which can be shaped through experience In addition mechanisms exist that affect neural integration time such as active conductances membrane oscillations 2439 and postinhibitory rebound 25 which can modify networklevel interactions Neural selectivity to temporal parameters of auditory stimuli has been studied extensively in echolocating bats These include selectivity to sound duration to delay between pulseecho sound pairs and to temporal rates of sound sequences all parameters that vary in the vocal signals produced by foraging bats One attribute of echolocation calls is signal duration a characteristic that changes markedly as bats approach a prey item Figure 1b Temporal ltering for sound duration has been reported at different levels of the auditory pathway and in several bat species 2627 and other mammals mouse 28 The underlying mech anism for durationtuned response pro les has been 39 39 using I 39 39 39 29 and intracellular recording methods 26 The response type appears to be rst created at the inferior colliculus 1C and is a consequence of the timing between excitatory and inhibitory converging inputs More recently by Current Opinion in Neurobiology 2003 13751 758 wwwcurrent opinioncom Figure 2 Neurobiology of echolocation in bats Moss and Sinha 753 Excitatory H a Auditory b Anterior cortical elds 7 Cingu39ate i cortex i Medial i geniculate v i Superior body 39 j colliculus Inferior Periaqueductal colliculus i gray i i l g Paralemniscal j j tegmental area Nuclei of the i i lateral lemniscus 1 Q i parabrachial i Dorsal i l nucleus i l j i Intermediate 1 l i i i l l j Nucleus Ventral r retroamlguus l i i 7 ii Nucleus ambiguus Key GABAergic Glycinergic Unidenti ed neurotransmitter n r n n r 4 Current Opinion in Neurobloiogy Major connections of the a ascending auditory system and descending auditory corticofugal projections The dashed box demarcates nuclei of he superior olivary complex 00 that includes the lateral superior olive the medial superior olive and the medial nucleus works Excitatory projections are shown with black lines inhibitory GABAergic with red lines inhibitory glycinergic projections are shown t b Selected projections of vocal production circuitry from cited projections are shown of the trapezoid body with green lines and connections with as yet unidentified neurotransmitters are shown with dashed black lines Abbreviations ON cochlear nucleus employing a two tone stimulus paradigm that used a robe tone at the neurons best frequency BF and best duration BD and a masking competing tone with a nonexcitatory NE duration the latency duration and decay of the afferent input inhibition was delineated 31 By manipulating the onset overlap and offset of the probe and masking tone the time course of inhibition has been shown to shape BD the duration tuning char acteristics and rst spike latency The distance between the bat and sonar target may be represented by the activity pro le in a population of neurons that respond selectively to two sounds whic simulate sonar cry and echo separated by a limited and biologically relevant range of temporal delays 3233 These delaytuned neurons are present in the midbrain 34 36 thalamus and cortex 3738 Delaytuned neu rons which are likely to be established at the level of the midbrain 3940 are sensitive to additional stimulus dimensions for example the absolute amplitude the spectral content and the temporal rate at which a series of stimulus pairs are presented Thus these neurons may not only encode target distance but also might potentially encode other stimulus dimensions 41 A recent study explored this by using two overlapping echoes temporally offset to simulate sonar reflections wwwcurrent opinioncom Current Opinion in Neurobiology 2003 137Si 758 754 Neurobiology of behaviour Figure 3 U50 3 Center 3 26 m E E a w 0 P PE E 200 Msec 39 lt gtlt 77 650 Center 26 a E E a w 0 P PE E 200 Msec lt gtlt gt 0 Contra Contra Spikesdata 5e 5 t N w o o f y g gg quot 32 s quot s 24 16 04306 w o 8 D Em delay mseC Azrrnutn degrees is o Spikesdata set N o Monopolar SC stimulation mecy mmv Frcqllml r kHz v 5 E E Q Bipolar SC stimulation Q PAG stimulation Fz unn and in A 15 m m a n an r m m N M su Spontaneous t mmnmm Q in a u All Current Oprnron rn Neurobiology a Neural recordings from the bat superior colliculus Echo delay tuned neuron in the bat SC shows a facilitated response to a pulse echo pair separated by 12 msec The response is vigorous at an current levels are shown in the u er rlght periaqueductal gray SC superior colliculus Adapted from 66 aztmuth of 26 deg contralateral to the recording site but falls off at 39 deg echo P pulse PE pulse and echo b Spatial response profiles of two SC neu c Sonar vocalizations elicited by electrical stimulation of the SC 1 and 2 and communication calls elicited by stimulation of the PAS Abbreviations rons that show selectivity to azimuth and delay Adapted from 36 3 Stimulation corner of each example Spontaneous vocalizations recorded from flying bats 4 Abbreviations PAG from closely spaced surfaces 42 The authors showed evidence for enhanced responses when pairs of partially overlapping echoes were presented after a simulated sonar cry suggesting a response pro le sensitive to both the temporal and spectral structure of stimuli Echolocating bats use sonar returns to localize objects in azimuth elevation and distance leading to the prediction that auditory neurons show spatial selectivity in 3D space A population of auditory neurons in the intermedi ate and deep superior colliculus SC of the bat Epm im fusms show 3D spatial response pro les 36 Figure 3a In this population of 3D neurons echo delaytuning is tagged to the azimuth and elevation of a sound source The representation offsD target location in the SC of the bat would be important for the coordination of sensory Current Opinion in Neurobiology 2003 13751 758 wwwcurrent opinioncom and motor signals that drive its acoustic orientation as changes in the bat s echolocation behavior are closely tied to position 15 The principal cue bats and other mammals use to localize the direction of high frequency sound is interaural level difference ILD Neurons that are excited by stimula tion of one ear and inhibited by stimulation of the other ear binaural response pro le created with excitatory inhibitory contralateralipsilateral inputs EI are thought to encode ILD This response type serves as the putative mechanism for spatial localization among bats the timing of convergent excitatory and inhibitory inputs onto these neurons being crucial in shaping their response patterns Timing is also important when one considers the duration over which the excitation or inhibition lasts as this can signi cantly affect the response of an ILD neuron to subsequent sounds This last point has recently been investigated 4339 by combining extracellular recordings with iontophoretic application of antagonists andor pre sentation of a pair of binaural sounds The persistent inhibition initiated at the dorsal nucleus of the lateral lemniscus DNLL prevents the DNLL from responding for a period of time As many 1C cells receive inhibitory input from DNLL they are temporarily transformed from strongly inhibited El neurons to weakly inhibited E1 or even monaural cells when DNLL activity is shut down Two clear implications arise both related to the temporal aspects of neural processing of multiple sounds The rst is that El response properties can change over time on the basis of the temporal pattern of binaural stimulation showing a state dependent response This is a consequence of timing and persistence of inhibition in this network of neurons and suggests that codes for spatial localization that are formed on the basis of El neuronal responses change with multiple stimuli The second implication is a consequence of the loss of the El property When 1C inhibitory input is reduced the El cells lose their ILD speci city and respond to sounds from a larger in region in space Recently there has been a growing interest in the proces sing of acoustic communication signals in the central nervous system CNS of echolocating bats Interestingly auditory regions traditionally studied in t e context of biosonarprocessing appear to play a role in communication signal processing Researchers nd that neural responses to communication signals depend on the temporal spectral characteristics of sounds similar to ndings for biosonar signals These results suggest that the process ing of sounds used for orientation and i 4 occurs through overlapping auditory networks 44 46 Corticofugal modulation Timedependent changes in basic auditory neuronal receptive eld parameters have been demonstrated 47 and recently expanded on in a series of elegant Neurobiology of echolocation in bats Moss and Sinha 755 experiments The research not only suggests the involve ment of the amygdala and cholinergic basal forebrain in auditory plasticity 48 but also strongly supports a role for corticofugal descending projections from the cortex modulation in adjusting neuronal receptive eld proper ties based on the salience of an auditory stimulus The effects have been observed at the level of auditory cortical elds ACF medial geniculate body MGB 1C 4950 and the cochlea 51 Using repetitive acoustic stimula tion fear conditioning focal electrical microstimulation of the primary auditory cortex Al or 1C or electrical microstimulation combined with auditory stimulation frequency response areas of auditory neurons in bats can be shi ted in an experiencedependent manner Neurons with temporal combinationselectivity are also influenced delaytuned 5253 durationtuned 54 at both cortical and subcortical levels he observed changes arise over the course of minutes and can last from a few seconds to hours 55 Protocols that involve associative learning eg electrical foot stimulation paired with a conditioning tone stimulus demonstrate changes in the receptive elds of AI neurons which can last up to 26h 56 The timecourse and underlying mechanism of this longterm change in best frequency as evaluated using Nmethyl Daspartate NMDA ago nists and blockers suggests the involvement of experi encedependent transcriptionally mediated processes in order to maintain longterm changes in a neuron s best frequency response area 57 59 The experiencedepen dent plasticity is observed in both FM 5660 and CF FM emitting bat species 5361 Work with similar pro tocols suggests that these corticofugal modulations are likely to generalize to other mammals 62 The implica tion in our view is that central and peripheral auditory processing in bats can be rapidly modulated to adjust the analysis of auditory signals in the context of changing vocal patterns and corresponding echoes The rapid experiencedependent plasticity clearly demonstrates that the inputs to neurons putatively involved in process ing 39 i i f i can be modu lated which permits a shift in their classical receptive eld arrangement Although these experiments do not employ echolocation behavior they do involve processing in neuronal populations sensitive to sonar calls and provide evidence for the potential range of plasticity possible when the bat is actively engaged in echolocation Motor production The behavioral context within which bats echolocate plays a central role in shaping the parameters of sonar 394 39 Sonar vucali aLiun produced by bats dur ing insect pursuit show considerable variation in duration an width spectral content and temporal patterning Figure 1 The speciesspeci c variations in call design relative motion ofthe bat with respect to its target changes in call structure and temporal patterning all influence the information available to the bat s auditory system wwwcurrent opinioncom Current Opinion in Neurobiology 2003 13751 758 756 Neurobiology of behaviour Using electrical and chemical microstimulation techni ques several brainstem regions involved in sonar vocal production have been studied Experiments have focused on midbrain regions identifying cytoarchitectural regions that elicit vocalizations and their interconnections In the midbrain the paralemniscal tegmentum area PLa peri aqueductal gray PAG reticular formation parabrachial nucleus PB SC and their direct or indirect projections to brainstem nucleus retroambiguus RA and nucleus ambiguus NA have been elucidated 63 656639 Researchers have proposed that sonar vocal production has evolved from vocal communication pathways 67 Recent work 6839 has identi ed dual pathways for the production of sonar and communication calls at the mid brain level in the neotropical FM bat P ylloslomu dis 010 One pathwa encompassing medial loci in the ventral PAG elicits classes of communication calls fol lowing chemical stimulation using dialysis of nonlesion concentrations of kainic acid A second pathway involves a locus in the lateral PAG and the PLa both eliciting sonar vocalizations when electrically and chemically sti mulated This experiment is the rst to show a dual role for the PAC a region considered to be mandatory in the vocal production pathway for communication calls 69 Experiments that investigate audio vocal interactions in the midbrain PLa show that the PLa can contribute to timing aspects of sonar vocalizations however lesions of the PLa do not eliminate the ability to produce sonar vocalization 70 which suggests that it is not a mandatory component of sonar vocal circuitry Recent experiments studied the role of the PB in sonar vocalizations in the CF FM bat R imap u femmequi um 71 Using ionto phoretic application of yamino butyric acid GABAergic an Lglutamate agonists and antagonists the authors demonstrated that the PB plays a role in the control of call frequency Application of muscimol GABAA agonist or CN X 6cyano7nitroquinoxaline Z3dione a gluta matergic anatagonist lowered the call frequency emitted at rest and during DSC behavior Conversely excitation induced by application of ocamino 3 hydroxy5methyl 4isoxazole propionic acid AMPA or by blocking inhibi tion using BMI bicuculline methiodide a GABAA antagonist increased sonar call frequencies The PB study of the horseshoe bat is the rst to demon strate midbrain control over production of sound fre quency 71 Stimulation experiments in other midbrain sites have only reported influences on the timing and number of sonar vocalizations but not speci cally the spectral content Sitedependent spectral characteristics of electrically elicited vocalizations have been reported for the anterior cingulate cortex ACC in another CF FM bat species Plemmluspdmellii 72 Electrical micro stimulation of the ACC elicited sonar vocalizations in rostral ACC whereas microstimulation of more caudal ACC regions elicited communication sounds In the case of PB microstimulation 71 control of the amount of excitation and inhibition more crucially impacts the frequency in the emitted CF component of the sonar vocalization Adaptive behaviors to dynamic stimuli require the inte gration of sensory information with motor programs to guide appropriate responses and awealth ofdata suggests that the midbrain SC plays a role in sensorimotor integra tion In individual species the functional organization of the SC reflects the importance of a particular sensory modality to an animal s goaldirected orienting responses n bats control of vocal signals is an integral part of its acoustic orienting system It is therefore not surprising that microstimulation of the bat SC elicits head and pinna movements along with the production of sonar vocaliza tions Figure 3b 66 73 Consistent with this result premotor bursts are recorded from the bat SC before each sonar vocalization 74 Combined these studies suggest that a complex circuit of interconnected nuclei serve to control the timing and spectrotemporal parameters of sonar vocalizations Most of these regions receive projections from auditory nuclei and they may have evolved from areas involved in the production of communication calls 6772 Conclusions As the echolocating bat orients in the environment and pursues insect prey its sound production patterns adapt to changing acoustic information 2315 These adap tive vocal production patterns provide a window to the information sought and collected by the active sonar system for acoustic imaging under different task condi tions Researchers have taken advantage of the signals recorded from echolocating bats engaged in behavioral tasks to employ biologically relevant stimuli for studies of auditory information processing in the brain The dynamic patterning ofthe bat s sonar vocalizations estab lishes a time scale for studying echo processing both at the level of individual sounds and across sequences of sounds Researchers have also begun to study premotor areas that are involved in sonar production A complete understanding of the neurobiology of bat echolocation however requires detailed CNS studies of the bat actively engaging in vocal motor behaviors that result in sonar echo returns Acknowledgements We are grateful for grant support from the National Institutes of Mental Health R01 MH56366 the Whitehall Foundation 2000e08e03eREN S9720 National Science Foundation IBNe0111973 te CF Mess and an Institutional National Research Service Award SeT32eDC00046 te SR Sinha We also acknowledge suppett of an NIH p30 gtsht te the centet of Comparative and Evolutionary Biology of Hearing R Dueling PI In addition we thank A Perez and W Xian for technical assistance and M Ayteltlh and K Ghose for comments en an earlier draft of the manuscript Current Opinion in Neurobiology 2003 13751 758 wwwcurrent opinioncom References and recommended reading Papers of particular interest published within the annual period of 39 39 39 d s reVIew have been highlighte a of special interest quot of outstanding interest 1 Moss CF Schnitzler H U Behavioral studies of auditory information processing In Hearing y Bats Springer Handbook of Auditory Research Edited by Fay RR Popper AN Berlin Springer Verlag 199587 145 Iquot Griffin D Listening in the Dark New Haven Yale University Press 1958 3 Valentine DE Moss CF Sensorimotor integration in bat sonar In Bats Phylogeny Morphology Echolocation and Conservation Biology Edited by Kunz TH Racey A Washington DC Smithsonian Institution Press 1998220 230 4 Jones KE Purvis A MacLarnon A Bininda Emonds OR Simmons NB A phylogenetic supertree of the bats Mammalia Chiroptera Biol Rev Camb Philos Soc 2002 77223 259 Squot Schnitzler HU Moss CF Denzinger A From spatial orientation to food acquisition in echolocating bats Trends Ecol Evol 2003 18386 394 57 Aldridge HDJN Rautenbach IL Morphology echolocation and resource partitioning in insectivorous bats J Anim Ecol 1987 56763 778 N Fenton MB Natural history and biosonar signals In Hearing by Bats Edited by Popper AN Fay RR New York Springer Verlag 199537 86 9 Neuweiler G Auditory adaptations for prey capture in echolocating bats Physiological Review 1990 70615 641 0 Schnitzler H U Kalko EKV Echolocation by insect eating bats BioScience 2001 51557 569 0 Schnitzler HU Die Ultraschall Ortungslaute der Hufeisen Fledermause Chiroptera Rhinolophidae in Verschiedenen Orientierungssituationen D39r nslation The ultrasound orientation sounds of the horseshoe bat Chiroptera Rhilophidae in different orientation situations Zeitschrift fuquotr Vergl Physiol 1968 57376 408 Metzner W Zhang S Smotherman M Doppler shift nsation behavior in horseshoe bats revisited auditory feedback controls both a crease an 39ncrease in call frequency J Exp Biol 2002 2051607 1616 Schnitzler H U Menne D Kober R Heblich K The acoustical image of fluttering insects in echolocating ats In Neuroethology and Behavioral Physiology Edited by Huber F Markel H Heidelberg Springer Verlag 1983 235 249 Iquot 9 Neuweiler G Evolutionary aspects of bat echolocation Journal of Comparative Physiology A 2003 189245 256 P Simmons JA The resolution of target range by echolocating bats J Acoust Soc Am 1973 54157 173 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leading inhibition versus onset evoked excitation the BD of a neuron can be determined 32 Feng AS Simmons JA Kick SA Echo detection and target ranging neurons in the auditory system of the bat Eptesicus fuscus Science 1978 202645 648 33 O Neill WE Suga N Target range sensitive neurons in the auditory cortex of the mustache bat Science 1979 20369 73 34 Dear SP Suga N Delay tuned neurons in the midbrain of the big brown bat J Neurophysiol 1995 731084 1100 35 Mittmann DH Wenstrup JJ Combination sensitive neurons in the inferior colliculus Hear Res 1995 90185 191 36 Valentine DE Moss CF Spatially selective auditory responses in the superior col 758 Neurobiology of behaviour Thomas JT Moss CF Vater M Eds Advances in the Study of Echolocation in Bats and Dolphins Chicago University of Chicago Press 2003 4 N Sanderson Ml Simmons JA Selectivity for echo spectral interference and delay in the auditory cortex of the big brown bat Eptesicus fuscus J Neurophysiol 2002 872823 2834 4 w Burger RM Pollak GD Reversible inactivation of the dorsal o nucleus of the lateral lemniscus revea s its role in t e processing of multiple sound sources in the inferior colliculus of bats J Neurosci 2001 21 4830 4843 The authors use extracellular recording with iontophoretic injections to stud quot 39 quot 39 mm e 39 39 39 in different locations in space They show by inactivating the DNLL that the El 5 u L properties of L the unenuv f rmed anew In addition they show that signals excitatory to El cells in IC result in inhibition of the contralateral DNLL lasting 10s of milliseconds T e long asting inhibition in e rives the l inhibition from DNLL which changes the response profiles of El cells in Ice essentially resulting in broader El tuning 44 Esser KH Condon OJ Suga N Kanwal JS Syntax processing by au itory cortical neurons in t e FM FM area of the mustached bat Pteronotus parnellii Proc Natl Acad Sci USA 1997 9414019 14024 4 w Bauer EE Klug A Pollak GD Spectral determination of r u L the 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Suga N Modulation of responses and frequency tuning of thalamic and collicular neurons by cortical activation in mustached bats J Neurophysiol 2000 84325 333 w P Ma X Suga N Corticofugal modulation of duration tuned neurons in the midbrain auditory nucleus in bats Proc NatlAcaol Sci USA 2001 9814060 14065 55 Zhou X Jen PH Brief and short term corticofugal modulation of subcortical auditory responses in the big brown bat Eptesicus fuscus J Neurophysiol 2000 843083 3087 01 SP Gao E Suga N Experience dependent plasticity in the auditory 1L u A system Proc Natl Acad Sci USA 2000 978081 8086 w i Tanahashi A HorikawaJ Suga N NMDA mediated facilitation in t e echo de ay tuned areas of the auditory co ex of the mustached bat Hear Res 1997 110219 228 01 9 Jen PH Feng RB Bicuculline application affects discharge pattern and pulse duration tuning characteristics of bat inferior collicular neurons JComp PhysiolA1999 184185 194 59 Schafe GE Nader K Blair HT LeDouxJE Memory consolidation 0 Pa ovian 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Afferent and efferent connections of the motor nuc eus of the laryngeal nerves J Comp PhysioA 1986 159689 699 66 Valentine DE Sinha SR Moss CF Orienting responses and o voca iza ions produced by microstimulation in the superior colliculus of the echolocating bat Eptesicus fuscus J Comp P ysiolANeuroethol Sens Neural Behav Physio2002 18 89 108 The authors report that microstimulation oft e at SC elicits head and pinna movements as observed in other mammals In addition SC A 4 A well as the reception of echolocation signals 67 Fenton MB Echolocation implications for ecology and evolution of bats Q Rev Biol 1983 5933 53 68 Fenzl T Schl39iller G Periaqueductal gray and the region of the o paralemni c have different functions in the control of vocalization in the neotropical bat Phyllostomus discolor Eur eurosci2002 161974 1986 The authors identify separate pathways for vocal production of commu nication sounds an 39 39 39 iontophoretic dialysis methods the sh 39 39 ocal communication soun s 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