BACKGROUND： Electrophysiological studies using brain slices have revealed that the developmental regulation of synaptic plasticity in vocal learning pathway is essential for song learning in zebra finches. Publicatio...BACKGROUND： Electrophysiological studies using brain slices have revealed that the developmental regulation of synaptic plasticity in vocal learning pathway is essential for song learning in zebra finches. Publications reporting in vivo electrophysiological investigation are scarce. Many aspects of neural mechanisms underlying song learning and production still remain uncertain. OBJECTIVE： To observe the efficacy of paired pulses and the effect on synaptic transmission induced by low-frequency stimulations, high-frequency stimulations, and theta-burst stimulations. DESIGN, TIME AND SETTING： A comparative observation. The experiment was conducted from October 2006 to October 2007 in the Neurophysiology Laboratory of South-China Normal University. MATERIALS： Twenty-four adult male zebra finches were supplied by the Department of Animal Experiment of College of Life Sciences, South China Normal University. A SEN-7203 stimulator （NIHON KOHDEN）, as well as a DSJ-731WF microelectrode amplifier and DSJ-F amplifier （provided by South-China Normal University）, were used to stimulate and record, respectively. METHODS： Animals were randomly divided into low-frequency, high-frequency, and theta-burst frequency stimulation groups. After recording evoked potentials, an input-output curve was evaluated. Subsequently, the efficacy of paired pulses with different stimulus intensity （1/3, 1/2, 2/3, or 3/4 of the value that induced the largest synaptic response）, as well as interpulse intervals （50, 75, and 100 ms）, was measured in each group. The test stimulus intensity was set to a level that evoked 1/2 or 1/3 amplitude of the maximum response. MAIN OUTCOME MEASURES： Changes in amplitude, slope, and area of evoked potentials elicited by different stimulus patterns. RESULTS： （1） Efficacy of paired pulses： there was significant paired-pulse facilitation in the high vocal center and robust nucleus of the arcopallium （HVC-RA） synapse. Efficacy decreased when paired-pulse intervals or stimulus intensities were increased. （2） Low-frequency stimulations at 1 Hz had a negligible effect on efficacy of synaptic transmission, while 5 Hz depressed synaptic transmission for only 5 minutes. （3） High-frequency stimulations at 50 Hz or 100 Hz induced synaptic depression that lasted for up to 30 minutes. （4） Theta-burst stimulation depressed synaptic transmission efficiency significantly for about 10 minutes. However, in contrast to low-frequency or high-frequency stimulations, theta-burst stimulations also induced slight potentiation of synaptic transmission for up to 60 minutes following depression phase, although the slope or area did not change. CONCLUSION： Paired pulses induced a remarkable efficacy of paired pulses in the high vocal center and robust nucleus of the arcopallium pathway. Low-frequency, high-frequency, or theta-burst frequency stimulation did not induce long-lasting changes in synaptic transmission.展开更多
BACKGROUND： Electrophysiological properties of the song nucleus have been revealed using conventional techniques, such as intracellular and extracellular recording. Research concerning the neuronal activation propert...BACKGROUND： Electrophysiological properties of the song nucleus have been revealed using conventional techniques, such as intracellular and extracellular recording. Research concerning the neuronal activation properties and regulations of the song system at the cellular and ion channel level may help reveal the neural mechanism of song learning. OBJECTIVE： To perform whole-cell recording of robust nucleus of the arcopallium （RA） neurons in brain slices from adult zebra finches （Taeniopygia guttata） and observe the action potential, sodium/potassium current and the spontaneous postsynaptic current of RA neurons. DESIGN, TIME AND SETTING： Self-controlled, neuroelectrophysiological experiment. The study was performed at the Neurophysiology Laboratory of South China Normal University from April to September 2008. MATERIALS： Flaming/Brown puller P-97 was purchased from Sutter Ins, USA; Axopatch 700B amplifier and Digidata 1332A converter were purchased from Axon Instrument, USA; pClamp software was provided by Axon Instrument, USA. METHODS： RA neurons were acutely isolated from 24 healthy male zebra finches. The action potential, voltage-gate sodium/potassium current and spontaneous postsynaptic current were recorded by whole-cell recording technology. Data were analyzed by pClamp software. MAIN OUTCOME MEASURES： The amplitude and frequency of the action potential, and the amplitude of the voltage-dependent and spontaneous postsynaptic currents, were measured. RESULTS： （1） Testing of action potential： Cells exhibited a stable current-voltage relationship following a series of hyperpolarization stepped currents, and an action potential was triggered by the spike threshold. All the recorded cells displayed repetitive firing following depolarizing current injection, with a frequency beyond 100 Hz. （2） Testing of voltage-gate currents： The inward and outward whole-cell currents were observed after a series of depolarizing voltage steps. The inward current disappeared following the application of tetrodotoxin and the outward current was significantly inhibited by application of 4-aminopyfidione and tetraethylammonium chloride. （3） Testing of spontaneous postsynaptic current： The majority of recorded cells exhibited an inward synaptic current when the membrane potential was maintained at -60 mV, with some cells exhibiting a robustly outward current when the membrane potential was maintained at -30 mV. Tetrodotoxin was unable to affect the spontaneous postsynaptic current. Following application of bicuculline [y-aminobutyric acid （A） receptor antagonist] and high concentration kynurenic acid （ionotropic glutamate receptor antagonist）, the inward and outward currents were completely inhibited. CONCLUSION： Under these experimental conditions, the action potential, sodium/potassium current and spontaneous postsynaptic current were recorded successfully in RA neurons. This indicates that the cells preserved relatively intact synaptic connections and normal physiological activity, which is required for investigating ion channels. The inward and outward whole-cell currents were sodium and potassium currents, respectively. The postsynaptic y-aminobutyric acid （A） receptors and ionotropic glutamate receptors contributed to the spontaneous postsynaptic current.展开更多
Objective To observe the sexual differences in electrophysiological properties of neurons in the robust nucleus of the arcopallium （RA） in adult zebra finches, and to provide the direct electrophysiological evidence...Objective To observe the sexual differences in electrophysiological properties of neurons in the robust nucleus of the arcopallium （RA） in adult zebra finches, and to provide the direct electrophysiological evidence for the sexual dimorphism of birdsong. Methods Whole-cell recording was used to record the spontaneous action potential firing rates from RA projection neurons in acute brain slices. Results The projection neurons of RA in male birds fired spontaneously at 10 Hz or above, while in female birds, the frequency was significantly lower, and even no firings could be detected. Conclusion There is a sexual difference in electrophysiological properties of projection neurons in RA, which may result from the difference in the levels of steroid hormones in birds.展开更多
Objective Motor control is encoded by neuronal activity. Small conductance Ca2＋-activated K＋ channels （SK channels） maintain the regularity and precision of firing by contributing to the afterhyperpolarization （...Objective Motor control is encoded by neuronal activity. Small conductance Ca2＋-activated K＋ channels （SK channels） maintain the regularity and precision of firing by contributing to the afterhyperpolarization （AHP） of the action potential in mammals. However, it is not clear how SK channels regulate the output of the vocal motor system in songbirds. The premotor robust nucleus of the arcopallium （RA） in the zebra finch is responsible for the output of song information. The temporal pattern of spike bursts in RA projection neurons is associated with the timing of the acoustic features of birdsong. Methods The firing properties of RA projection neurons were analyzed using patch clamp whole-cell and cell-attached recording techniques. Results SK channel blockade by apamin decreased the AHP amplitude and increased the evoked firing rate in RA projection neurons. It also caused reductions in the regularity and precision of firing. RA projection neurons displayed regular spontaneous action potentials, while apamin caused irregular spontaneous firing but had no effect on the firing rate. In the absence of synaptic inputs, RA projection neurons still had spontaneous firing, and apamin had an evident effect on the firing rate, but caused no significant change in the firing regularity, compared with apamin application in the presence of synaptic inputs. Conclusion SK channels contribute to the maintenance of firing regularity in RA projection neurons which requires synaptic activity, and consequently ensures the precision of song encoding.展开更多
Environmental stressors can negatively affect avian cognitive abilities, potentially reducing fitness, for example by altering response to predators, display to mates, or memory of locations of food. We expand on curr...Environmental stressors can negatively affect avian cognitive abilities, potentially reducing fitness, for example by altering response to predators, display to mates, or memory of locations of food. We expand on current knowledge by investigating the effects of dietary mercury, a ubiquitous en- vironmental pollutant and known neurotoxin, on avian cognition. Zebra finches Taeniopygia gut- tata were dosed for their entire lives with sub-lethal levels of mercury, at the environmentally rele- vant dose of 1.2 parts per million. In our first study, we compared the dosed birds with controls of the same age using tests of three cognitive abilities： spatial memory, inhibitory control, and color association. In the spatial memory assay, birds were tested on their ability to learn and remember the location of hidden food in their cage. The inhibitory control assay measured their ability to ig- nore visible but inaccessible food in favor of a learned behavior that provided the same reward. Finally, the color association task tested each bird＇s ability to associate a specific color with the presence of hidden food. Dietary mercury negatively affected spatial memory ability but not inhibi- tory control or color association. Our second study focused on three behavioral assays not tied to a specific skill or problem-solving： activity level, neophobia, and social dominance. Zebra finches exposed to dietary mercury throughout their lives were subordinate to, and more active than, con- trol birds. We found no evidence that mercury exposure influenced our metric of neophobia. Together, these results suggest that sub-lethal exposure to environmental mercury selectively harms neurological pathways that control different cognitive abilities, with complex effects on be- havior and fitness.展开更多
鸣禽高级发声中枢(high vocal center,HVC)至弓状皮质栎核(robust nucleus of the arcopallium,RA)的突触传递是鸣唱运动通路中的关键部分。本文运用在体场电位电生理记录的方法,研究了成年雄性斑胸草雀(Taeniopygia guttata)HVC-RA突...鸣禽高级发声中枢(high vocal center,HVC)至弓状皮质栎核(robust nucleus of the arcopallium,RA)的突触传递是鸣唱运动通路中的关键部分。本文运用在体场电位电生理记录的方法,研究了成年雄性斑胸草雀(Taeniopygia guttata)HVC-RA突触的电生理特性。实验结果显示,刺激HVC,在RA内所记录到的诱发场电位幅度较小。配对脉冲检测发现,HVC-RA突触传递具有明显的配对脉冲易化特性。当以强直刺激作用于HVC,RA内诱发场电位随即显著减小,并在15min内逐渐恢复,表明HVC-RA突触传递在强直刺激过后出现了短时抑制。该通路的突触传递特性可能与其在发声控制中的作用有关。以上的实验结果为进一步研究发声运动过程中的突触可塑性提供了资料。展开更多
斑胸草雀是一种研究鸣禽发声学习的模式动物,其鸣唱核团体积具有明显的性别差异,主要与体内雄激素水平的差异有关.以成年雌性斑胸草雀为研究对象,通过埋植睾酮,人为提高雌鸟体内雄激素水平,利用冰冻切片技术,结合尼氏染色法观察埋植前后...斑胸草雀是一种研究鸣禽发声学习的模式动物,其鸣唱核团体积具有明显的性别差异,主要与体内雄激素水平的差异有关.以成年雌性斑胸草雀为研究对象,通过埋植睾酮,人为提高雌鸟体内雄激素水平,利用冰冻切片技术,结合尼氏染色法观察埋植前后HVC(high vocal center)、RA(robust nucleus of the arcopallium)核团体积的变化.实验分为雌鸟埋植组、正常雌鸟组与正常雄鸟组.实验结果表明:正常雄鸟的RA、HVC核团体积均显著大于正常雌鸟;雌鸟埋植睾酮后,RA核团体积显著增加,神经元数目未发生明显改变,HVC核团体积显著增加,神经元数目显著增加.综上,埋植睾酮后能使HVC、RA核团体积趋于雄性化.展开更多
中脑丘间复合体背内侧核(dorsomedial nucleus of the intercollicular complex,DM)是鸣禽鸟的基本发声中枢.应用电声理学与声谱分析相结合的方法研究鸣禽斑胸草雀(Taeniopygia guttata))DM核团对其鸣声的调控模式.结果显示:刺激斑胸草...中脑丘间复合体背内侧核(dorsomedial nucleus of the intercollicular complex,DM)是鸣禽鸟的基本发声中枢.应用电声理学与声谱分析相结合的方法研究鸣禽斑胸草雀(Taeniopygia guttata))DM核团对其鸣声的调控模式.结果显示:刺激斑胸草雀左、右侧DM产生的诱发叫声声谱图类似,不具有典型谐音频带,刺激右侧DM核团产生的诱发叫声主能量区带是刺激左侧的2.4倍,声长为1.3倍,提示右侧DM核团对诱发叫声的影响较大.损毁左、右侧DM后自鸣声中产生大量恒频声,这是在正常鸣叫声中没有的现象,提示DM核团对鸣叫声的频率调制起着至关重要的作用.损毁右侧DM后自鸣声声长为左侧的1.3倍,主能量区带为1.3倍,提示右侧损毁DM对自鸣声的影响较大.左右两侧差异体现斑胸草雀的DM核团对于非习得性鸣声的调控具有右侧优势.展开更多
基金the National Natural Science Foundation of China, No. 30370197,30570232
文摘BACKGROUND： Electrophysiological studies using brain slices have revealed that the developmental regulation of synaptic plasticity in vocal learning pathway is essential for song learning in zebra finches. Publications reporting in vivo electrophysiological investigation are scarce. Many aspects of neural mechanisms underlying song learning and production still remain uncertain. OBJECTIVE： To observe the efficacy of paired pulses and the effect on synaptic transmission induced by low-frequency stimulations, high-frequency stimulations, and theta-burst stimulations. DESIGN, TIME AND SETTING： A comparative observation. The experiment was conducted from October 2006 to October 2007 in the Neurophysiology Laboratory of South-China Normal University. MATERIALS： Twenty-four adult male zebra finches were supplied by the Department of Animal Experiment of College of Life Sciences, South China Normal University. A SEN-7203 stimulator （NIHON KOHDEN）, as well as a DSJ-731WF microelectrode amplifier and DSJ-F amplifier （provided by South-China Normal University）, were used to stimulate and record, respectively. METHODS： Animals were randomly divided into low-frequency, high-frequency, and theta-burst frequency stimulation groups. After recording evoked potentials, an input-output curve was evaluated. Subsequently, the efficacy of paired pulses with different stimulus intensity （1/3, 1/2, 2/3, or 3/4 of the value that induced the largest synaptic response）, as well as interpulse intervals （50, 75, and 100 ms）, was measured in each group. The test stimulus intensity was set to a level that evoked 1/2 or 1/3 amplitude of the maximum response. MAIN OUTCOME MEASURES： Changes in amplitude, slope, and area of evoked potentials elicited by different stimulus patterns. RESULTS： （1） Efficacy of paired pulses： there was significant paired-pulse facilitation in the high vocal center and robust nucleus of the arcopallium （HVC-RA） synapse. Efficacy decreased when paired-pulse intervals or stimulus intensities were increased. （2） Low-frequency stimulations at 1 Hz had a negligible effect on efficacy of synaptic transmission, while 5 Hz depressed synaptic transmission for only 5 minutes. （3） High-frequency stimulations at 50 Hz or 100 Hz induced synaptic depression that lasted for up to 30 minutes. （4） Theta-burst stimulation depressed synaptic transmission efficiency significantly for about 10 minutes. However, in contrast to low-frequency or high-frequency stimulations, theta-burst stimulations also induced slight potentiation of synaptic transmission for up to 60 minutes following depression phase, although the slope or area did not change. CONCLUSION： Paired pulses induced a remarkable efficacy of paired pulses in the high vocal center and robust nucleus of the arcopallium pathway. Low-frequency, high-frequency, or theta-burst frequency stimulation did not induce long-lasting changes in synaptic transmission.
基金the National Natural Science Foundation of China,No.30570232the Natural Science Foundation of Guangdong Province,No. 05005910Key Laboratory of Ecology and Environmental Science in Guangdong Higher Education
文摘BACKGROUND： Electrophysiological properties of the song nucleus have been revealed using conventional techniques, such as intracellular and extracellular recording. Research concerning the neuronal activation properties and regulations of the song system at the cellular and ion channel level may help reveal the neural mechanism of song learning. OBJECTIVE： To perform whole-cell recording of robust nucleus of the arcopallium （RA） neurons in brain slices from adult zebra finches （Taeniopygia guttata） and observe the action potential, sodium/potassium current and the spontaneous postsynaptic current of RA neurons. DESIGN, TIME AND SETTING： Self-controlled, neuroelectrophysiological experiment. The study was performed at the Neurophysiology Laboratory of South China Normal University from April to September 2008. MATERIALS： Flaming/Brown puller P-97 was purchased from Sutter Ins, USA; Axopatch 700B amplifier and Digidata 1332A converter were purchased from Axon Instrument, USA; pClamp software was provided by Axon Instrument, USA. METHODS： RA neurons were acutely isolated from 24 healthy male zebra finches. The action potential, voltage-gate sodium/potassium current and spontaneous postsynaptic current were recorded by whole-cell recording technology. Data were analyzed by pClamp software. MAIN OUTCOME MEASURES： The amplitude and frequency of the action potential, and the amplitude of the voltage-dependent and spontaneous postsynaptic currents, were measured. RESULTS： （1） Testing of action potential： Cells exhibited a stable current-voltage relationship following a series of hyperpolarization stepped currents, and an action potential was triggered by the spike threshold. All the recorded cells displayed repetitive firing following depolarizing current injection, with a frequency beyond 100 Hz. （2） Testing of voltage-gate currents： The inward and outward whole-cell currents were observed after a series of depolarizing voltage steps. The inward current disappeared following the application of tetrodotoxin and the outward current was significantly inhibited by application of 4-aminopyfidione and tetraethylammonium chloride. （3） Testing of spontaneous postsynaptic current： The majority of recorded cells exhibited an inward synaptic current when the membrane potential was maintained at -60 mV, with some cells exhibiting a robustly outward current when the membrane potential was maintained at -30 mV. Tetrodotoxin was unable to affect the spontaneous postsynaptic current. Following application of bicuculline [y-aminobutyric acid （A） receptor antagonist] and high concentration kynurenic acid （ionotropic glutamate receptor antagonist）, the inward and outward currents were completely inhibited. CONCLUSION： Under these experimental conditions, the action potential, sodium/potassium current and spontaneous postsynaptic current were recorded successfully in RA neurons. This indicates that the cells preserved relatively intact synaptic connections and normal physiological activity, which is required for investigating ion channels. The inward and outward whole-cell currents were sodium and potassium currents, respectively. The postsynaptic y-aminobutyric acid （A） receptors and ionotropic glutamate receptors contributed to the spontaneous postsynaptic current.
基金supported by the National Natural Science Foundation of China (No. 30970363,30570232)the Natural Science Foundation of Guangdong Province, China (No. 05005910)
文摘Objective To observe the sexual differences in electrophysiological properties of neurons in the robust nucleus of the arcopallium （RA） in adult zebra finches, and to provide the direct electrophysiological evidence for the sexual dimorphism of birdsong. Methods Whole-cell recording was used to record the spontaneous action potential firing rates from RA projection neurons in acute brain slices. Results The projection neurons of RA in male birds fired spontaneously at 10 Hz or above, while in female birds, the frequency was significantly lower, and even no firings could be detected. Conclusion There is a sexual difference in electrophysiological properties of projection neurons in RA, which may result from the difference in the levels of steroid hormones in birds.
基金supported by grants from the National Natural Science Foundation of China (30970363 and 31172092)
文摘Objective Motor control is encoded by neuronal activity. Small conductance Ca2＋-activated K＋ channels （SK channels） maintain the regularity and precision of firing by contributing to the afterhyperpolarization （AHP） of the action potential in mammals. However, it is not clear how SK channels regulate the output of the vocal motor system in songbirds. The premotor robust nucleus of the arcopallium （RA） in the zebra finch is responsible for the output of song information. The temporal pattern of spike bursts in RA projection neurons is associated with the timing of the acoustic features of birdsong. Methods The firing properties of RA projection neurons were analyzed using patch clamp whole-cell and cell-attached recording techniques. Results SK channel blockade by apamin decreased the AHP amplitude and increased the evoked firing rate in RA projection neurons. It also caused reductions in the regularity and precision of firing. RA projection neurons displayed regular spontaneous action potentials, while apamin caused irregular spontaneous firing but had no effect on the firing rate. In the absence of synaptic inputs, RA projection neurons still had spontaneous firing, and apamin had an evident effect on the firing rate, but caused no significant change in the firing regularity, compared with apamin application in the presence of synaptic inputs. Conclusion SK channels contribute to the maintenance of firing regularity in RA projection neurons which requires synaptic activity, and consequently ensures the precision of song encoding.
基金This work was funded by an award from the National Science Foundation [to J.P.S. and D.A.C.] （IOS1257590} and an English-Stonehouse fellowship [to J.P.S. and A.S.F.] awarded by the College of William and Mary.
文摘Environmental stressors can negatively affect avian cognitive abilities, potentially reducing fitness, for example by altering response to predators, display to mates, or memory of locations of food. We expand on current knowledge by investigating the effects of dietary mercury, a ubiquitous en- vironmental pollutant and known neurotoxin, on avian cognition. Zebra finches Taeniopygia gut- tata were dosed for their entire lives with sub-lethal levels of mercury, at the environmentally rele- vant dose of 1.2 parts per million. In our first study, we compared the dosed birds with controls of the same age using tests of three cognitive abilities： spatial memory, inhibitory control, and color association. In the spatial memory assay, birds were tested on their ability to learn and remember the location of hidden food in their cage. The inhibitory control assay measured their ability to ig- nore visible but inaccessible food in favor of a learned behavior that provided the same reward. Finally, the color association task tested each bird＇s ability to associate a specific color with the presence of hidden food. Dietary mercury negatively affected spatial memory ability but not inhibi- tory control or color association. Our second study focused on three behavioral assays not tied to a specific skill or problem-solving： activity level, neophobia, and social dominance. Zebra finches exposed to dietary mercury throughout their lives were subordinate to, and more active than, con- trol birds. We found no evidence that mercury exposure influenced our metric of neophobia. Together, these results suggest that sub-lethal exposure to environmental mercury selectively harms neurological pathways that control different cognitive abilities, with complex effects on be- havior and fitness.
基金This work was supported by the National Natural Science Foundation of China(No.30370197,30570232)the Natural Science Foundation of Guangdong Province(No.05005910).
文摘鸣禽高级发声中枢(high vocal center,HVC)至弓状皮质栎核(robust nucleus of the arcopallium,RA)的突触传递是鸣唱运动通路中的关键部分。本文运用在体场电位电生理记录的方法,研究了成年雄性斑胸草雀(Taeniopygia guttata)HVC-RA突触的电生理特性。实验结果显示,刺激HVC,在RA内所记录到的诱发场电位幅度较小。配对脉冲检测发现,HVC-RA突触传递具有明显的配对脉冲易化特性。当以强直刺激作用于HVC,RA内诱发场电位随即显著减小,并在15min内逐渐恢复,表明HVC-RA突触传递在强直刺激过后出现了短时抑制。该通路的突触传递特性可能与其在发声控制中的作用有关。以上的实验结果为进一步研究发声运动过程中的突触可塑性提供了资料。
文摘斑胸草雀是一种研究鸣禽发声学习的模式动物,其鸣唱核团体积具有明显的性别差异,主要与体内雄激素水平的差异有关.以成年雌性斑胸草雀为研究对象,通过埋植睾酮,人为提高雌鸟体内雄激素水平,利用冰冻切片技术,结合尼氏染色法观察埋植前后HVC(high vocal center)、RA(robust nucleus of the arcopallium)核团体积的变化.实验分为雌鸟埋植组、正常雌鸟组与正常雄鸟组.实验结果表明:正常雄鸟的RA、HVC核团体积均显著大于正常雌鸟;雌鸟埋植睾酮后,RA核团体积显著增加,神经元数目未发生明显改变,HVC核团体积显著增加,神经元数目显著增加.综上,埋植睾酮后能使HVC、RA核团体积趋于雄性化.
文摘中脑丘间复合体背内侧核(dorsomedial nucleus of the intercollicular complex,DM)是鸣禽鸟的基本发声中枢.应用电声理学与声谱分析相结合的方法研究鸣禽斑胸草雀(Taeniopygia guttata))DM核团对其鸣声的调控模式.结果显示:刺激斑胸草雀左、右侧DM产生的诱发叫声声谱图类似,不具有典型谐音频带,刺激右侧DM核团产生的诱发叫声主能量区带是刺激左侧的2.4倍,声长为1.3倍,提示右侧DM核团对诱发叫声的影响较大.损毁左、右侧DM后自鸣声中产生大量恒频声,这是在正常鸣叫声中没有的现象,提示DM核团对鸣叫声的频率调制起着至关重要的作用.损毁右侧DM后自鸣声声长为左侧的1.3倍,主能量区带为1.3倍,提示右侧损毁DM对自鸣声的影响较大.左右两侧差异体现斑胸草雀的DM核团对于非习得性鸣声的调控具有右侧优势.
