猫外膝体细胞方位方向敏感性与时间空间频率的关系

在１２只成年猫上,用微电极记录了外膝体神经元对不同方位（方向）的移动正弦光栅刺激的反应,测定了细胞在不同时间、空间频率下的方位、方向调谐特性。在一定的时频范围内,外膝体神经元的方位敏感性受时间频率的影响不大,而方向敏感性受时间频率的影响相对较大：在其它参数不变的情况下,改变时间频率可产生相当不同的方向调谐特性。外膝体神经元一般在近于截止频率的较高空间频率下具有较强的方位敏感性,而在近于最优空间频率的较低空间频率下呈现出较强的方向敏感性。多数方位、方向敏感性细胞对空间频率的依赖较大,即只在一个较窄的空间频率范围内呈现出它们的敏感性或随空间频率的不同敏感性强度产生较大的变化;也有少数方位、方向敏感性细胞在较宽的空间频率范围内呈现明显的方位、方向敏感性。结果提示,外膝体神经元的方位敏感性和方向敏感性的产生,有着不同的时间、空间机制     

尾核阿片μ受体参与电针及皮层SmⅠ区对束旁核伤害性反应的抑制

为探索尾核（ｃａｕｄａｔｅｎｕｃｌｅｕｓ,Ｃｄ）是否参与电针及皮层体感运动Ⅰ区（ｓｅｎｓｏｒｉｍｏｔｏｒａｒｅａⅠｏｆｔｈｅｃｅｒｅｂｒａｌｃｏｒｔｅｘ,ＳｍⅠ）对束旁核（ｐａｒａｆａｓｃｉｃｕｌａｒｎｕｃｌｅｕｓ,Ｐｆ）神经元伤害性反应的调节,以及Ｃｄ中阿片受体是否参与并通过何种受体参与这一调节,本实验用Ｃｄ头部化学毁损及微量注射阿片受体拮抗剂的方法,观察到Ｃｄ毁损前电针及兴奋皮层均可抑制Ｐｆ的伤害性反应,而毁损后这种抑制效应消失;注射纳洛酮或阿片μ受体拮抗剂βＦＮＡ后,电针及兴奋皮层ＳｍⅠ区对Ｐｆ伤害性反应的抑制作用被取消,而分别注射δ和κ受体拮抗剂ＩＣＩ１７４,８６４和ｎｏｒＢＮＩ则不产生影响。基于已证明大脑皮层参与电针对Ｐｆ伤害性反应的调节,本结果提示：Ｃｄ参与针刺镇痛中皮层ＳｍⅠ区对Ｐｆ神经元伤害性反应的抑制,Ｃｄ中阿片肽主要通过μ受体参与抑制作用。     

刺激中缝背核调制小脑间位核神经元电活动

刺激中缝背核（ｄｏｒｓａｌｒａｐｈｅｎｕｃｌｅｕｓ,ＤＲ）可以引起小脑间位核（ｉｎｔｅｒｐｏｓｅｄｎｕｃｌｅｕｓ,ＩＮ）神经元抑制,兴奋和双相（抑制－兴奋和兴奋－抑制）３种不同类型的反应,其中以抑制反应为主（７６．０％）,多数细胞的反应潜伏期〈３０ｍｓ。ＩＮ细胞的自发放电频率为５－１２０Ｈｚ,自发放电频率高的神经元群体对ＤＲ刺激的反应率却比自发放电频率低的群体低。静脉注射５－ＨＴ２／１ｃ受体阻断剂     

刺激中缝背核对大鼠小脑皮层下核团神经元电活动的影响

我们首次观察了电刺激大鼠中缝背核（ＤＲ）对小脑核团（ＤＣＮ）──内侧核（ＭＮ）,间位核（ＩＮ）和外侧核（ＬＮ）神经元电活动的影响。结果表明：刺激ＤＲ可引起ＤＣＮ神经元的抑制、兴奋和双相（兴奋－抑制、抑制－兴奋）三种不同类型的反应,其中以抑制性反应为主（７６％－９０％）;反应的潜伏期为１０—８４ｍｓ,但大多数细胞呈现小于３０ｍｓ的短潜伏期反应;ＤＣＮ细胞的自发放电频率为５－１２０Ｈｚ,自发放电频率高的神经元群体对ＤＲ刺激的反应比率却比自发放电频率低的群体低;静脉注射５－ＨＴ２／１ｃ受体阻断剂ｍｅｔｈｙｓｅｒｇｉｄｅ可以阻断ＤＣＮ细胞对ＤＲ刺激的抑制性反应（６６．７％－８３．３％）。这些结果提示中缝－小脑５－ＨＴ能纤维传入系统可能通过对ＤＣＮ细胞电活动的调制作用参予小脑的感觉运动整合过程。     

猫皮层SⅠ区神经元对刺激VPL的反应及其膜电学特性的研究

本文采用细胞内记录技术,研究了猫皮层第一躯体感觉区（ｐｒｉｍａｒｙｓｏｍａｔｏｓｅｎｓｏｒｙｃｏｒｔｅｘａｒｅａ,ＳⅠ区）躯体伤害感受神经元膜的电学特性和对刺激腹后外侧核（ｖｅｎｔｒａｌｐｏｓｔｅｒｉｏｒｌａｔｅｒａｌｎｕｃｌｅｕｓ,ＶＰＬ核）的反应。极化电流绝对值小于或等于１０ｎＡ时,伤害感受神经元ＩＶ极相关（ｒ＝０９６）,整流作用不明显;极化电流绝对值大于１０ｎＡ时,在两个方向上发生整流,ＩＶ曲线表现为Ｓ型,其中伤害感受神经元的整流作用较非伤害感受神经元明显。伤害性感受神经元Ｒｍ、τ、Ｃｍ明显大于非伤害感受神经元（Ｐ＜００１或Ｐ＜００５）。刺激ＶＰＬ与刺激隐神经在ＳⅠ区伤害感受神经元的诱发反应中存有相似与不同两种形式。用细胞内电位记录方法证明了单一神经元有会聚现象。结果提示,ＳⅠ区伤害感受神经元与非伤害感受神经元可能在细胞膜形态结构、细胞体积大小等方面存在有意义的差别,从而反映其不同的生理功能。     

猫后内侧上雪氏区视神经元对运动棒反应的取向和方向特征

猫后内侧上雪区（ｐｏｓｔｅｒｏｍｅｄｉａｌｌａｔｅｒａｌｓｕｐｒａｓｙｌｖｉａｎａｒｅａ,ＰＭＬＳ）的绝大多数神经元（１７１／２００）对运动棒的取向调谐,６２％（１２４／２００）细胞的取向调谐宽度（半高波宽）小于９０°：按方向选择性和取向选择性可分辨出几类特征明显的细胞类型：１、强取向和强方向选择性细胞;２、强取向调谐的双向选择细胞;３、弱取向调谐的强方向选择细胞;４、无取向无方向选择性细胞;以及５、特征不明显的或中间类型细胞。它们与最近光学记录揭示的鹰猴中颞叶视区（ｍｉｄｄｌｅｔｅｍｐｏｒａｌｖｉｓｕａｌａｒｅａ,ＭＴ）的组织有很好的吻合。     

听神经元的频率调谐

频率作为声音的一个重要参数,在听敏感神经元对声音进行分析和编码过程中扮演重要角色。一般用频率调谐曲线来表示听敏感神经元的频率调谐特性,并用Qn(10,30,50)值表达频率调谐曲线的尖锐程度,Qn值越大,频率调谐曲线也越尖锐,神经元的频率调谐能力越好,对频率的分辨能力越高。从听觉外周到中枢,听敏感神经元的频率调谐逐级锐化,而这种锐化主要是由听中枢的多种抑制性神经递质的作用而产生的,其中起主要作用的是GABA能和甘氨酸能神经递质。此外,离皮层调控,双侧下丘间的联合投射以及弱噪声前掩蔽等因素也会影响听敏感神经元的频率调谐特性。     

N-甲基-D-天冬氨酸(NMDA)及荷包牡丹碱对不同年龄蝙蝠下丘神经元听反应的影响

分别对出生后第２周和第４周的中华鼠耳蝠（Ｍｙｏｔｉｓｃｈｉｎｅｎｓｉｓ）中脑下丘中央核１２１个听神经元进行了考察,同时与成年动物进行比较。结果表明,微电泳ＮＭＤＡ对下丘中央核绝大部分听神经元（２周龄、４周龄和成年动物分别为９６％、９５％和９６％）具有易化性影响,表现为听反应脉冲发放率增加、反应阈值下降、频率调谐曲线增宽。ＮＭＤＡ的易化性效应部分地与动物的周龄相关,表现为对听反应阈值和听反应脉冲发放率的易化效应在出生后２－４周逐渐增强,但４周以后的易化效应趋缓;微电泳ＧＡＢＡＡ受体的特异性拮抗剂Ｂｉｃｕｃｕｌｉｎｅ,对神经元的听反应也呈现易化性影响,表现为听反应脉冲发放率增加,反应阈值降低和频率调谐曲线增宽等。实验结果还提示,ｂｉｃｕｃｕｌｉｎｅ对下丘神经元听反应的易化作用,在出生后早期呈持续增强趋势。ＮＭＤＡ和ｂｉｃｕｃｕｌｉｎｅ的效应与动物周龄之间的关系略有差异,是否提示着下丘中央核ＮＭＤＡ受体和ＧＡＢＡＡ受体在出生后的发育进程有所差别,尚待进一步探讨。     

猫皮层18区神经元整合野特性的研究

用双重正弦调制移动光栅图形研究了猫皮层１８区神经元整合野特性。同在１７区所观察到的结果一样,在１８区神经元传统感受野（ＣＲＦ）的外面,也存在着大范围的易化或抑制性整合野（ＩＦ）。抑制和易化性ＩＦ神经元分别占７０％和３０％。ＩＦ对刺激光栅的时间频率、空间频率、方位和方向都有选择性。对大多数细胞来说,ＩＦ的这些调谐特性都与其对应的ＣＲＦ的特性相似。以上结果提示,１８区神经元能够对ＣＲＦ和ＩＦ内的图形特征进行整合,这种整合在复杂图形识别中起重要作用。     

γ—氨基丁酸能抑制可镜化大棕蝠听皮层神经元频率调谐

本实验使用了9只成年健康的大棕蝠（Eptesicus fuscus）。采用双声刺激和多管电极电泳导入荷包牡丹碱（bicuculline,Bic）的方法,研究了γ－氨基丁酸（γ－aminobutyric,GABA）能抑制在锐化听皮层（primary auditory cortex,AC,即初级听皮层）神经元频率调谐中的作用。结果发现：正常AC神经元的频率调谐曲线表现出单峰开放式、多峰开放式和单峰封闭式2种类型;用双声刺激方法研究证实,至AC神经元的抑制性输入能被抑制性声刺激所激活,且这种神经抑制有自身的最佳频率,根据其对兴奋反应的影响程度和系统地改变抑制性声刺激的强度,可在兴奋性频率调谐曲线或兴奋区的高频边或／和低频边测出抑制性频率调谐曲线或抑制区;当这种抑制性输入被抑制性声刺激激活后,能降低阈上10fB声强引起的兴奋反应的发放率,抑制效率随抑制声刺激强度的增强而加强;电泳GABAa受体拮抗剂荷包牡丹碱Bic后,可不同程度地去GABA能抑制参与构成至AC神经元的抑制性输入,在正常情况下这种抑制有助于提高中枢听神经元的信号／噪声比和频率分析能力,并锐化频率调谐,因此本结果在正常情况下这种抑制有助于提高中枢听神经元的信号／噪声比和频率分析能力,并锐化频率调谐,因此本结果提示,声音的各参量中所包含的信息从外周传入中枢后,随着中枢的升级,逐级抽提整合成若干特征,直至在AC形成某种“声像（sound image)”,对大多数AC神经元而言,GABA能抑制在该过程中起关键作用。     

经验改变大鼠听皮层神经元的特征频率

应用常规电生理学技术,以神经元的特征频率和频率调谐曲线为指标,研究大鼠听皮层神经元特征频率的可塑性. 结果表明,在给予的条件刺激频率和神经元特征频率相差1.0 kHz范围内,条件刺激可诱导50%以上神经元特征频率发生完全偏移,并可分为向频率调谐曲线的低频端偏移、高频端偏移,或两侧均可偏移三种类型. 其中,神经元的特征频率高、Q₁₀-dB值大和频率调谐曲线对称指数大于零的神经元,其特征频率偏向频率调谐曲线高频端的概率更高. 结果提示,经验可改变大鼠听皮层神经元的特征频率,为深入研究中枢神经元功能活动可塑性的机制提供了重要实验资料.     

大鼠生后发育过程中听皮层神经元特征频率的可塑性

应用常规电生理学技术,以神经元的特征频率和频率调谐曲线为指标,分别在生后2、3、4、5、6和8周龄SD大鼠上,研究生后发育过程中,听皮层神经元特征频率的可塑性.结果表明,在给予条件刺激频率和神经元特征频率相差1.0kHz范围内,条件刺激都可诱导各年龄组神经元特征频率向频率调谐曲线的低频端、高频端或调谐曲线的两端相应的偏移.特征频率偏移的概率与年龄相关.随着年龄的增长,特征频率偏移的比例下降,而不偏移的比例则上升.随着年龄增长,那些Q10-dB值大和频率调谐曲线对称指数大于零的神经元,特征频率偏移到频率调谐曲线高频端的比例增加更为明显(P<0.01).诱导特征频率完全偏移的时程和特征频率恢复的时程也与动物的年龄相关,随着年龄增长,诱导和恢复时程都明显延长(P<0.05).结果提示,大鼠听皮层神经元特征频率的可塑性与生后年龄相关,为深入研究中枢神经元功能活动可塑性的机制提供了重要实验资料.     

电刺激大马蹄蝠听皮层对下丘神经元听觉敏感性的影响

实验在１２只大马蹄蝠上进行。用常规电生理学方法研究了电刺激听皮层对下丘２１２个神经元的听反应的影响,结果表明,有３２个神经元的听反应被抑制,１９个神经元的听反应褐易化。     

Responses of neurons in the rat auditory and associative cortices to tonal stimuli

Activity of single neurons and mass evoked potentials (EP) were recorded from the auditory (area 41) and associative (area 39) cortices in acute experiments on rats anesthetized with urethane, nembutal, or chloralose; pure tones were used as acoustic stimuli. The EP appearing in response to a wide range of sound tones on the surface of the auditory and associative cortices were dissimilar in their latency and shape. For neurons exhibiting stable responses, the frequency-threshold curves (FTC) were plotted.Weak and variable responses of neurons were observed under slight urethane anesthesia. Nembutal anesthesia increased the responsiveness of neurons and the probability of appearing of late components in the responses. Chloralose anesthesia was characterized by extension of frequency range perceived by a neuron, while its sharpness of tuning remained unchanged. Under all types of anesthesia employed, the responses recorded from the associative cortex neurons had longer latencies than those recorded from the auditory cortex neurons. Neurons exhibiting the frequency selectivity were much less numerous in the associative cortex than in the auditory cortex. The former neurons were often characterized by intermittent FTC and they responded to a more extended frequency range. No clear tonotopic organization was found in the associative cortex.Neirofiziologiya/Neurophysiology, Vol. 25, No. 5, pp. 343–349, September–October, 1993.     

锐化蝙蝠听皮层神经元频率调谐的柱特征

用双声刺激和多管电极方法在 6只大棕蝠 (bigbrownbat,Eptesicusfuscus)的 98个神经元上研究了锐化 (sharpening)蝙蝠听皮层 (primaryauditorycortex ,AC)神经元频率调谐的柱特征。结果发现 ,电极直插在 1个电极通道内连续记录到多个神经元时 ,它们锐化频率调谐的抑制性调谐曲线或抑制区基本相似。电极与AC表面呈 45°斜向推入使其跨越多个功能柱时 ,可观察到锐化频率调谐的抑制区构成也随电极进入不同的功能柱而发生相应的改变。两种不同的电极插入方式均证明锐化AC神经元频率调谐的神经抑制呈柱状组构。这些神经元组合起来排列在同一听觉功能柱内 ,构成AC频率分析的基本功能组构单位“微频率处理器”。实验中还观察到多峰频率调谐曲线神经元 ,它们在声通讯和声定位中不同波谱区域的时间匹配中起作用。此外 ,也有理由认为多峰调谐神经元亦被用于作为复杂波谱信息的“高级调谐预处理器” ,从而极大地提高了神经元对频率分析的能力。为研究锐化频率调谐的神经抑制机制 ,用多管电极电泳γ -氨基丁酸 (γ aminobutyricacid ,GA BA)能a受体拮抗剂荷包牡丹碱 (bicuculline ,Bic)至所记录的神经元 ,发现能大部分或几乎全部取消抑制区 ,从而表明在正常情况下GABA能抑制参与构成锐化AC神经元频率调谐的抑制区 ,     

Effects of GABA-mediated inhibition on direction-dependent frequency tuning in the frog inferior colliculus

Earlier studies from our laboratory have shown that the frequency selectivity of neurons in the frog inferior colliculus is direction dependent. The goal of this study was to test the hypotheses that gamma-aminobutyric acid or GABA (but not glycine)-mediated synaptic inhibition was responsible for the direction-dependence in frequency tuning, and that GABA acted through creation of binaural inhibition. We performed single unit recordings and investigated the unit's free-field frequency tuning, and/or the unit's response to the interaural level differences (under dichotic stimulation), before and during local applications of antagonists specific to gamma-aminobutyric acid a and glycine receptors. Our results showed that application of bicuculline produced a broadening of free-field frequency tuning, and differential changes in free-field frequency tuning depending on sound direction, i.e., more pronounced at azimuths at which the unit exhibited narrower frequency tuning under the pre-drug condition, thereby typically abolishing direction dependence in tuning. Application of strychnine produced no change in frequency tuning. The results from dichotic stimulation further revealed that bicuculline typically elevated and/or flattened the unit's interaural-level-difference response function, indicating a reduction in the strength of binaural inhibition. Our study provides evidence that gamma-aminobutyric acid-mediated binaural inhibition is important for direction dependence in frequency tuning. Accepted: 24 July 1998     

Lateral sharpening of cortical frequency tuning by approximately balanced inhibition

Cortical inhibition plays an important role in shaping neuronal processing. The underlying synaptic mechanisms remain controversial. Here, in vivo whole-cell recordings from neurons in the rat primary auditory cortex revealed that the frequency tuning curve of inhibitory input was broader than that of excitatory input. This results in relatively stronger inhibition in frequency domains flanking the preferred frequencies of the cell and a significant sharpening of the frequency tuning of membrane responses. The less selective inhibition can be attributed to a broader bandwidth and lower threshold of spike tonal receptive field of fast-spike inhibitory neurons than nearby excitatory neurons, although both types of neurons receive similar ranges of excitatory input and are organized into the same tonotopic map. Thus, the balance between excitation and inhibition is only approximate, and intracortical inhibition with high sensitivity and low selectivity can laterally sharpen the frequency tuning of neurons, ensuring their highly selective representation.     

Multiple mechanisms shape selectivity for FM sweep rate and direction in the pallid bat inferior colliculus and auditory cortex

The inferior colliculus and auditory cortex of the pallid bat contain a large percentage of neurons that are highly selective for the direction and rate of the downward frequency modulated (FM) sweep of the bat’s echolocation pulse. Approximately 25% of neurons tuned to the echolocation pulse respond exclusively to downward FM sweeps. This review focuses on the finding that this selectivity is generated by multiple mechanisms that may act alone or in concert. In the inferior colliculus, selectivity for sweep rate is shaped by at least three mechanisms: shortpass or bandpass tuning for signal duration, delayed high-frequency inhibition that prevents responses to slow sweep rates, and asymmetrical facilitation that occurs only when two tones are presented at appropriate delays. When acting alone, the three mechanisms can produce essentially identical rate selectivity. Direction selectivity can be produced by two mechanisms: an early low-frequency inhibition that prevents responses to upward sweeps, and the same asymmetrical two-tone inhibition that shapes rate tuning. All mechanisms except duration tuning are also present in the auditory cortex. Discussion centers on whether these mechanisms are redundant or complementary.     

大棕蝠下丘听神经元频率调谐的方向敏感性

实验在8只大棕蝠(Eptesicusfuscus)上进行.选取了5个声源方位记录部位对侧80°(C-80°)、40°(C-40°)、0°和同侧40°(I-40°)、80°(I-80°).下丘听神经元的频率调谐曲线的形状有三种V型、封闭型和U型.以C-40°的频率调谐曲线为对照,声源方位改变时,多数神经元频率调谐曲线的形状和最佳频率不变.多数神经元在声源方位由记录部位对侧转向同侧时,频率调谐曲线的锐度(以QndB值为指标)由小变大;神经元在I-80°、I-40°的QndB值显著高于C-80°、C-40°、0°的QndB值.在所观察的47个神经元中,5个声源方位的最低阈值的平均值有显著差异,由高到低的分布为MTI-80°＞MTI-40°＞MT0°＞MTc-80°＞MTc-40°.我们推测,声源方位对神经元频率调谐特性影响的机制,可能是抑制性和兴奋性输入的比例不同而相互整合的结果.