下丘神经元对不同声刺激模式的反应特性研究
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摘要
本研究在自由声场条件下,采用在体神经电生理记录方法,以大蹄蝠(leaf-nosed bat, Hipposideros armiger)、几内亚长翼蝠(Miniopterus magnater)和小鼠(mouse, Mus musculus, Km)为模型,研究了“下丘(inferior colliculus, IC)神经元对不同声刺激模式的反应特性”,获得了一系列有意义的研究结果：
1.所录制到的大蹄蝠回声定位叫声为恒频-调频(constant frequency-frequency modulation, CF-FM)信号,一般含有2-3个谐波,1-3谐波的CF频率依次为33.3(±0.2)、66.5(±0.3)、99.4(±0.5)kHz,其中第2谐波为其主频(dominant frequency, DF)。所记录大蹄蝠IC神经元的最佳频率(best frequency, BF)和记录深度之间存在线性相关(n=169,r=0.716,P<0.001),BF调谐在DF的神经元具有较低的最小阈值(minimal threshold, MT)和较高的Q值。当用CF和FM声单独刺激时,神经元对1次刺激只产生1个on反应(single-on response)。当用CF-FM声刺激时,大部分(76%)IC神经元仅对CF-FM声刺激的开始产生反应,即single-on反应,将其称之为single-on神经元；其余(24%)的IC神经元则对CF-FM声刺激的CF和FM成分的开始分别产生反应,即双反应(double-on response),将其称之为double-on神经元。实验观察到double-on神经元对单独的FM成分反应的阈值和潜伏期要比单独的CF成分更高和更长；double-on神经元对CF-FM刺激反应的潜伏期短于single-on神经元。这些结果表明,大蹄蝠IC神经元的频率调谐与回声定位信号的主频之间密切相关,BF位于主频及其附近的神经元其频率调谐和分析能力最强,CF-FM声信号中的FM成分在定型神经元的发放类型、阈值、反应潜伏期和放电率方面起着重要作用；并提示这两种反应类型的神经元有可能在蝙蝠巡航期间,分别扮演着根据回声定性靶物和获取靶物细节信息的角色。
2.大蹄蝠IC神经元对CF和CF-FM声刺激的强度放电率函数均表现为单调型、饱和型和非单调型,在CF-FM声刺激下,单调型神经元比例下降,饱和型和非单调型比例增加；且神经元的最佳强度(best amplitude, BA) (84.4±11.3 dB SPL)和动力学范围(dynamic range, DR)(22.5±8.5 dB)均要比CF刺激下的BA(93.0±9.8 dB SPL)(P<0.01)和DR(27.5±9.4 dB)(P<0.05)低；而强度放电率函数曲线的斜率(slope) (4.1±1.7%/dB)则要高于CF刺激(3.1±1.4%/dB)(P<0.05)。进一步分析不同强度下FM成分对神经元放电率的影响,结果显示,在低强度(MT+30 dB)下则降低了CF的兴奋作用(P<0.001)。基于以上结果推测,低强度下FM中邻近CF的频率成分发挥了CF的持续效应；而高强度下,FM中偏离CF较远的频率成分激活了低频边高阈值的侧抑制神经元,从而抑制了神经元的放电率。结果提示CF-FM蝙蝠回声定位信号FM成分能够提高了下丘神经元的声强敏感性。
3.由于回声频率和强度均可发生改变,而时程则不然,成为蝙蝠识别自身声脉冲的“标签”。本实验通过向几内亚长翼蝠给不同的刺激呈现率(presentation rate, PR)的单声刺激和不同间隔的脉冲-回声对刺激,来研究IC神经元在不同PR条件下的时程调谐以及对声刺激下的前掩蔽。结果显示,随着PR的增加,IC神经元的时程调谐类型由长通或者全通型向带通、短通型转变,大多数神经元关键带(critical band)带宽变窄(P<0.001)；前掩蔽的效果与神经元的时程调谐特性无关,R值随着时程的增加而增加(P<0.001)。这些结果表明,PR增加导致带通和短通型神经元比例增加,使得神经元跟随高脉冲重复率能力增强,其结果正好与蝙蝠在回声定位过程中不断增加发声脉冲重复率和缩短时程的回声定位行为相匹配,而前掩蔽效应与声刺激时程的正相关有利于蝙蝠在捕食过程中更多地获取回声信息。
4.用对声刺激模式在几内亚长翼蝠的IC所获得研究结果显示,当掩蔽效应指数“R”下降50%时,高、低频边的半带宽(half-band width),可将受前掩蔽影响的神经元分为低频边长效掩蔽型、高频边长效掩蔽型和双边等效或均衡掩蔽型；所有神经元的高频边半带宽(half-band widthhigh)与低频边半带宽(half-band widthlow)之间存在线性相关(n=24,r=0.47,P<0.05)；50%神经元的half-band widthlow显著大于half-band widthhigh(P<0.001),25%神经元half-band widthhigh显著大于half-band width,low (P<0.05),其余25%的神经元的half-band widthhigh与half-band widthlow之间无显著差异(P>0.05)。基于这些结果推测,由偏离BF声刺激所产生的强掩蔽效应,或许能为蝙蝠的发声抑制理论和有回声环境中的听觉抑制效应提供某种实验证据。
5.在对声刺激模式下,研究了小鼠IC神经元前掩蔽所致的膜电位变化,结果显示,神经元对单声刺激诱发的动作电位(action potential, AP)发放主要有单AP(n=11)、双AP(n=8)和串AP(n=5)发放3种模式；所诱发的抑制性突触后电位(inhibitory postsynaptic potential, IPSP)可出现在兴奋前或/和兴奋后。58.3%的神经元在对声间隔大于掩蔽声诱发的兴奋后IPSP时程时,探测声诱发的AP恢复到单声刺激水平；41.7%的神经元在对声间隔小于或等于掩蔽声诱发的兴奋后IPSP时程时,AP发放恢复到单声刺激水平；而几乎所有的神经元(23/24,95.8%)只有对声间隔大于掩蔽声诱发的兴奋后IPSP时程时,探测声反应潜伏期才完全恢复到单声刺激水平。但发放数完全恢复所需的平均对声间隔与掩蔽声诱发的兴奋后IPSP时程之间并无显著差异(P>0.05),而潜伏期恢复所需的平均对声间隔显著大于掩蔽声诱发的兴奋后IPSP寸程(P<0.05)。这些结果提示突触后抑制和神经元的膜电位状态等参与了前掩蔽的形成。
Under free field stimulation condition, we studied the auditory response properties of inferior collicular (IC) neurons of leaf-nosed bat (Hipposideros armiger), Miniopterus magnater, and mouse (Mus musculus) to different sound stimulus patterns using in vivo neuroelectrophysiological recording, and obtained the results as following.
1. The recorded echolocation calls of the leaf-nosed bats were a constant frequency-frequency modulation (CF-FM) signal with 2-3 harmonics. The recoded CF components of three harmonics from 18 leaf-nosed bats (n=18) were 33.3±0.2 kHz,66.5±0.3 kHz, and 99.4±0.5 kHz, respectively, and the second one among these three harmonics was its dominant frequency. The best frequencies (BFs) had linear correlation with recording depth of all the recorded IC neurons (n=169, r=0.716, P<0.001). The neurons tuned at DF had lower minimal threshold (MT) and larger Qn values than other neurons. All the neurons could generate a single onset response (single-on response, SO) to both CF and FM sound stimuli. When using CF-FM sound stimulus, most (76%) neurons were still to generate a SO response to CF-FM stimulus, which were called SO responders, but the remaining (24%) neurons could generate severally an onset response (double-on (DO) response) to both CF and FM components of CF-FM sound, which were called DO responders. On the other hand, the DO responders have higher minimum threshold (MT) and longer latency to the alone FM stimulus than alone CF. The results showed that there was certain relationship between echolocation call and frequency tuning of IC neurons in the leaf-nosed bats, and the FM component of the CF-FM sound took part in shapping discharge pattern, latency and firing rate of IC neurons. The data also suggested that these two types of responders might play respectively different roles in echo analysis during hunting.
2. Under CF and CF-FM stimulation conditions, rate-amplitude functions (RAF) of the leaf-nosed bats'IC neurons could be classified into three types, i.e monotonic, saturated, and non-monotonic. CF-FM stimulus induced higher proportions of the nonmonotonic and saturated than CF stimulus. Comparison of best amplitude (BA), dynamic range (DR), and slope of the RAEs obtained by CF and CF-FM stimuli, CF-FM stimulus decreased BA from 93.0±9.8 to 84.4±11.3 dB SPL (P<0.01), DR from 27.5±9.4 to 22.5±8.5 dB (P<0.05), and increased slope from 3.1±1.4 to 4.1±1.7%/dB (P<0.05). On the other hand, the result showed that FM component could increase firing rates of the IC neurons at less than 30 dB above MT while could decrease firing rates at more than 30 dB above MT (P<0.001). It is suggested that the mechanism underlying the firing rate increasing at lower amplitude could be a summational effect induced by partial frequencies of FM near CF, while the firing rate decreasing at higher amplitude was perhaps because of lateral inhibition activated by those frequencies of FM away from the CF at higher amplitude. Therefore, FM of the echolocation call of CF-FM bat could increase the sensitivity of IC neurons to sound amplitude.
3. Generally, the frequency and amplitude of echo would change during echolocation, but the echo duration could keep no change, as such the echo duration was served as a "tag" to recognize a bat's own call. By presenting single sound stimulus of different presentation rate (PR) and pulse-echo pair stimulus of different duration to the Miniopterus magnater, we examined the duration tuning and forward masking of IC neurons. The duration tuning of most IC neurons were changed from all-pass or long-pass toward band-pass or short-pass and their critical bandwidths were decreased (P<0.001) with PR increasing. There was no any relationship between types of duration tuning and forward masking, the R value of forward masking increased with the masker's duration increasing for all types of duration tuning (P<0.001). These results indicated that the higher PR could induce a large proportion of short-pass and band-pass neurons and enhance the ability of the IC neurons to follow high PR. So, it was matched better with the echolocation of bats which increase their emitted pulse repletion rate with approaching target during hunting.
4. The forward masking of IC neurongs in Miniopterus magnater decreased gradually with off-BF of masker frequency toward low and high frequency sides under pulse-echo pair stimulation condition. The neurons affected by maskers could be categorized into three types, i.e. low frequency side long-masking, high frequency side long-masking, and both side equal-masking neurons according to their half-band widths of low (half-band widthlow) and high (half-band widthhigh) frequency sides and the forward masking index "R" at 50% forward masking. There was linear correlated between half-band widths of low and high frequency sides in all neurons (n=24, r=0.47, P<0.05). Among theses neurons, half-band widthlow in 50% of the neurons was remarkable more than half-band widthhigh (P<0.001), half-band widthhigh, in 25% of the neurons was remarkable more than half-band widthlow (P<0.05), and half-band widths of low and high frequency sides in the remaining 25% of neurons were equal (P>0.05). These results suggested that forward masking caused by off-BF sound could be the base of depression induced by bat vocalization and precedence effect in the hearing.
5. Under paired sound stimulation condition, the changes in membrane potentials of mouse IC neurons were studied. The types of action potentials (AP) elicited by single sound could be classified into the single (n=11), double (n=8) and train (n=5) while inhibitory postsynaptic potential (IPSP) evoked by sound stimuation could occur at pre- or/and post-AP firing. The AP firing to probe was completely recovery when the paired sound gap was longer than the duration of masker-evoked IPSP at post-AP in 58.3% of IC neurons. The AP firing of the remaining (41.7%) IC neuronms could be completely recovery at paired sound gap less than or equal to the duration of masker-evoked IPSP at post-AP. However, the response latency to probe could be completely recovery when paired sound gap was longer than the duration of masker-evoked IPSP at post-AP in almost all the IC neurons (23/24,95.8%). By statistical analysis, only latency recovery need paired sound gap longer than the duration of the masker-evoked IPSP at post-AP (P<0.05). These results suggested that the postsynaptic inhibition and membrane potential level of the IC neurons might participate in the forward masking.
1.所录制到的大蹄蝠回声定位叫声为恒频-调频(constant frequency-frequency modulation, CF-FM)信号,一般含有2-3个谐波,1-3谐波的CF频率依次为33.3(±0.2)、66.5(±0.3)、99.4(±0.5)kHz,其中第2谐波为其主频(dominant frequency, DF)。所记录大蹄蝠IC神经元的最佳频率(best frequency, BF)和记录深度之间存在线性相关(n=169,r=0.716,P<0.001),BF调谐在DF的神经元具有较低的最小阈值(minimal threshold, MT)和较高的Q值。当用CF和FM声单独刺激时,神经元对1次刺激只产生1个on反应(single-on response)。当用CF-FM声刺激时,大部分(76%)IC神经元仅对CF-FM声刺激的开始产生反应,即single-on反应,将其称之为single-on神经元；其余(24%)的IC神经元则对CF-FM声刺激的CF和FM成分的开始分别产生反应,即双反应(double-on response),将其称之为double-on神经元。实验观察到double-on神经元对单独的FM成分反应的阈值和潜伏期要比单独的CF成分更高和更长；double-on神经元对CF-FM刺激反应的潜伏期短于single-on神经元。这些结果表明,大蹄蝠IC神经元的频率调谐与回声定位信号的主频之间密切相关,BF位于主频及其附近的神经元其频率调谐和分析能力最强,CF-FM声信号中的FM成分在定型神经元的发放类型、阈值、反应潜伏期和放电率方面起着重要作用；并提示这两种反应类型的神经元有可能在蝙蝠巡航期间,分别扮演着根据回声定性靶物和获取靶物细节信息的角色。
2.大蹄蝠IC神经元对CF和CF-FM声刺激的强度放电率函数均表现为单调型、饱和型和非单调型,在CF-FM声刺激下,单调型神经元比例下降,饱和型和非单调型比例增加；且神经元的最佳强度(best amplitude, BA) (84.4±11.3 dB SPL)和动力学范围(dynamic range, DR)(22.5±8.5 dB)均要比CF刺激下的BA(93.0±9.8 dB SPL)(P<0.01)和DR(27.5±9.4 dB)(P<0.05)低；而强度放电率函数曲线的斜率(slope) (4.1±1.7%/dB)则要高于CF刺激(3.1±1.4%/dB)(P<0.05)。进一步分析不同强度下FM成分对神经元放电率的影响,结果显示,在低强度(
3.由于回声频率和强度均可发生改变,而时程则不然,成为蝙蝠识别自身声脉冲的“标签”。本实验通过向几内亚长翼蝠给不同的刺激呈现率(presentation rate, PR)的单声刺激和不同间隔的脉冲-回声对刺激,来研究IC神经元在不同PR条件下的时程调谐以及对声刺激下的前掩蔽。结果显示,随着PR的增加,IC神经元的时程调谐类型由长通或者全通型向带通、短通型转变,大多数神经元关键带(critical band)带宽变窄(P<0.001)；前掩蔽的效果与神经元的时程调谐特性无关,R值随着时程的增加而增加(P<0.001)。这些结果表明,PR增加导致带通和短通型神经元比例增加,使得神经元跟随高脉冲重复率能力增强,其结果正好与蝙蝠在回声定位过程中不断增加发声脉冲重复率和缩短时程的回声定位行为相匹配,而前掩蔽效应与声刺激时程的正相关有利于蝙蝠在捕食过程中更多地获取回声信息。
4.用对声刺激模式在几内亚长翼蝠的IC所获得研究结果显示,当掩蔽效应指数“R”下降50%时,高、低频边的半带宽(half-band width),可将受前掩蔽影响的神经元分为低频边长效掩蔽型、高频边长效掩蔽型和双边等效或均衡掩蔽型；所有神经元的高频边半带宽(half-band widthhigh)与低频边半带宽(half-band widthlow)之间存在线性相关(n=24,r=0.47,P<0.05)；50%神经元的half-band widthlow显著大于half-band widthhigh(P<0.001),25%神经元half-band widthhigh显著大于half-band width,low (P<0.05),其余25%的神经元的half-band widthhigh与half-band widthlow之间无显著差异(P>0.05)。基于这些结果推测,由偏离BF声刺激所产生的强掩蔽效应,或许能为蝙蝠的发声抑制理论和有回声环境中的听觉抑制效应提供某种实验证据。
5.在对声刺激模式下,研究了小鼠IC神经元前掩蔽所致的膜电位变化,结果显示,神经元对单声刺激诱发的动作电位(action potential, AP)发放主要有单AP(n=11)、双AP(n=8)和串AP(n=5)发放3种模式；所诱发的抑制性突触后电位(inhibitory postsynaptic potential, IPSP)可出现在兴奋前或/和兴奋后。58.3%的神经元在对声间隔大于掩蔽声诱发的兴奋后IPSP时程时,探测声诱发的AP恢复到单声刺激水平；41.7%的神经元在对声间隔小于或等于掩蔽声诱发的兴奋后IPSP时程时,AP发放恢复到单声刺激水平；而几乎所有的神经元(23/24,95.8%)只有对声间隔大于掩蔽声诱发的兴奋后IPSP时程时,探测声反应潜伏期才完全恢复到单声刺激水平。但发放数完全恢复所需的平均对声间隔与掩蔽声诱发的兴奋后IPSP时程之间并无显著差异(P>0.05),而潜伏期恢复所需的平均对声间隔显著大于掩蔽声诱发的兴奋后IPSP寸程(P<0.05)。这些结果提示突触后抑制和神经元的膜电位状态等参与了前掩蔽的形成。
Under free field stimulation condition, we studied the auditory response properties of inferior collicular (IC) neurons of leaf-nosed bat (Hipposideros armiger), Miniopterus magnater, and mouse (Mus musculus) to different sound stimulus patterns using in vivo neuroelectrophysiological recording, and obtained the results as following.
1. The recorded echolocation calls of the leaf-nosed bats were a constant frequency-frequency modulation (CF-FM) signal with 2-3 harmonics. The recoded CF components of three harmonics from 18 leaf-nosed bats (n=18) were 33.3±0.2 kHz,66.5±0.3 kHz, and 99.4±0.5 kHz, respectively, and the second one among these three harmonics was its dominant frequency. The best frequencies (BFs) had linear correlation with recording depth of all the recorded IC neurons (n=169, r=0.716, P<0.001). The neurons tuned at DF had lower minimal threshold (MT) and larger Qn values than other neurons. All the neurons could generate a single onset response (single-on response, SO) to both CF and FM sound stimuli. When using CF-FM sound stimulus, most (76%) neurons were still to generate a SO response to CF-FM stimulus, which were called SO responders, but the remaining (24%) neurons could generate severally an onset response (double-on (DO) response) to both CF and FM components of CF-FM sound, which were called DO responders. On the other hand, the DO responders have higher minimum threshold (MT) and longer latency to the alone FM stimulus than alone CF. The results showed that there was certain relationship between echolocation call and frequency tuning of IC neurons in the leaf-nosed bats, and the FM component of the CF-FM sound took part in shapping discharge pattern, latency and firing rate of IC neurons. The data also suggested that these two types of responders might play respectively different roles in echo analysis during hunting.
2. Under CF and CF-FM stimulation conditions, rate-amplitude functions (RAF) of the leaf-nosed bats'IC neurons could be classified into three types, i.e monotonic, saturated, and non-monotonic. CF-FM stimulus induced higher proportions of the nonmonotonic and saturated than CF stimulus. Comparison of best amplitude (BA), dynamic range (DR), and slope of the RAEs obtained by CF and CF-FM stimuli, CF-FM stimulus decreased BA from 93.0±9.8 to 84.4±11.3 dB SPL (P<0.01), DR from 27.5±9.4 to 22.5±8.5 dB (P<0.05), and increased slope from 3.1±1.4 to 4.1±1.7%/dB (P<0.05). On the other hand, the result showed that FM component could increase firing rates of the IC neurons at less than 30 dB above MT while could decrease firing rates at more than 30 dB above MT (P<0.001). It is suggested that the mechanism underlying the firing rate increasing at lower amplitude could be a summational effect induced by partial frequencies of FM near CF, while the firing rate decreasing at higher amplitude was perhaps because of lateral inhibition activated by those frequencies of FM away from the CF at higher amplitude. Therefore, FM of the echolocation call of CF-FM bat could increase the sensitivity of IC neurons to sound amplitude.
3. Generally, the frequency and amplitude of echo would change during echolocation, but the echo duration could keep no change, as such the echo duration was served as a "tag" to recognize a bat's own call. By presenting single sound stimulus of different presentation rate (PR) and pulse-echo pair stimulus of different duration to the Miniopterus magnater, we examined the duration tuning and forward masking of IC neurons. The duration tuning of most IC neurons were changed from all-pass or long-pass toward band-pass or short-pass and their critical bandwidths were decreased (P<0.001) with PR increasing. There was no any relationship between types of duration tuning and forward masking, the R value of forward masking increased with the masker's duration increasing for all types of duration tuning (P<0.001). These results indicated that the higher PR could induce a large proportion of short-pass and band-pass neurons and enhance the ability of the IC neurons to follow high PR. So, it was matched better with the echolocation of bats which increase their emitted pulse repletion rate with approaching target during hunting.
4. The forward masking of IC neurongs in Miniopterus magnater decreased gradually with off-BF of masker frequency toward low and high frequency sides under pulse-echo pair stimulation condition. The neurons affected by maskers could be categorized into three types, i.e. low frequency side long-masking, high frequency side long-masking, and both side equal-masking neurons according to their half-band widths of low (half-band widthlow) and high (half-band widthhigh) frequency sides and the forward masking index "R" at 50% forward masking. There was linear correlated between half-band widths of low and high frequency sides in all neurons (n=24, r=0.47, P<0.05). Among theses neurons, half-band widthlow in 50% of the neurons was remarkable more than half-band widthhigh (P<0.001), half-band widthhigh, in 25% of the neurons was remarkable more than half-band widthlow (P<0.05), and half-band widths of low and high frequency sides in the remaining 25% of neurons were equal (P>0.05). These results suggested that forward masking caused by off-BF sound could be the base of depression induced by bat vocalization and precedence effect in the hearing.
5. Under paired sound stimulation condition, the changes in membrane potentials of mouse IC neurons were studied. The types of action potentials (AP) elicited by single sound could be classified into the single (n=11), double (n=8) and train (n=5) while inhibitory postsynaptic potential (IPSP) evoked by sound stimuation could occur at pre- or/and post-AP firing. The AP firing to probe was completely recovery when the paired sound gap was longer than the duration of masker-evoked IPSP at post-AP in 58.3% of IC neurons. The AP firing of the remaining (41.7%) IC neuronms could be completely recovery at paired sound gap less than or equal to the duration of masker-evoked IPSP at post-AP. However, the response latency to probe could be completely recovery when paired sound gap was longer than the duration of masker-evoked IPSP at post-AP in almost all the IC neurons (23/24,95.8%). By statistical analysis, only latency recovery need paired sound gap longer than the duration of the masker-evoked IPSP at post-AP (P<0.05). These results suggested that the postsynaptic inhibition and membrane potential level of the IC neurons might participate in the forward masking.
引文
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