Hendrickson A E, Van Brederode J F, Mulligan K A, Celio M R
Department of Biological Structure, University of Washington, Seattle 98195.
J Comp Neurol. 1991 May 22;307(4):626-46. doi: 10.1002/cne.903070409.
The development of immunoreactivity for the calcium-binding proteins parvalbumin (PV) and calbindin-D28K (Cal) was studied in Macaca nemestrina striate cortex from fetal (F) 60 days to postnatal (P) 5 + years. We correlated changes in PV and Cal staining patterns with the well-documented developmental sequence for primate striate cortex neuron generation and maturation, synaptogenesis, and thalamocortical axon interactions in an attempt to deduce a functional role for these proteins. Our major findings is that Cal and PV have diametrically opposed developmental patterns except in layer 1. At F60 days both are present only in neurons of layer 1 and the number of labeled cell bodies and processes increases up to F125 days. Almost all Cal+ and PV+ cells in layer 1 disappear by P12 weeks. Cal is present by F113 days in pyramidal and stellate neurons, particularly layers 4-6. The numbers and staining density of cells in layers 2-6 increases up to birth and then both decline by P9-12 weeks. Supragranular layers show a second increase in Cal labeling from P20-36 weeks, and then there is a slow decline to the adult pattern which is reached by P1-2 years. Cell bodies in layers 4A, 4C alpha, and deep 4C beta are heavily Cal+ during pre- and early post-natal periods, but upper 4C beta remains unlabeled. PV is not seen until F155-162 days in layers 2-6. Large stellate and a few pyramidal cells appear first in layers 5/6 and 4C alpha, but PV+ stellate neurons are found in all layers except 4C beta by P6 weeks. Layer 4C beta contains a few PV+ cell bodies at P3 weeks, and light neuropile staining at P6 weeks, but then PV labeling rapidly increases so that by P12 weeks the density of 4C beta exceeds that of 4C alpha. Striate cortex has an adult pattern of cell number and neuropile density by P20 weeks. These developmental patterns suggest that the highest density of Cal cell body staining does not correlate with synaptogenesis, or the postnatal critical period of visually driven, binocular interactions. Rather Cal appears when lateral geniculate axons arrive in cortex, persists over the entire span of thalamocortical interactions, and disappears during the decline of cortical plasticity. The appearance of PV is highly correlated with the onset of complex visually driven activity at birth, while both the number of PV+ cell bodies and the density of PV+ neuropile reach adult levels coincident with the completion of thalamocortical connections.
在猕猴(Macaca nemestrina)的纹状皮层中,研究了从胎儿60天到出生后5年以上钙结合蛋白小白蛋白(PV)和钙结合蛋白-D28K(Cal)免疫反应性的发育情况。我们将PV和Cal染色模式的变化与灵长类纹状皮层神经元生成和成熟、突触形成以及丘脑皮质轴突相互作用的详细发育序列相关联,试图推断这些蛋白质的功能作用。我们的主要发现是,除了第1层外,Cal和PV具有完全相反的发育模式。在胎儿60天时,两者仅存在于第1层的神经元中,标记的细胞体和突起数量在胎儿125天时增加。到出生后12周时,第1层中几乎所有的Cal+和PV+细胞都消失了。Cal在胎儿113天时出现在锥体细胞和星状神经元中,特别是第4 - 6层。第2 - 6层中的细胞数量和染色密度在出生前增加,然后在出生后9 - 12周时下降。颗粒上层在出生后20 - 36周时Cal标记出现第二次增加,然后缓慢下降至出生后1 - 2年达到的成年模式。在出生前和出生后早期,第4A层、4Cα层和深层4Cβ层中的细胞体Cal+染色很重,但上层4Cβ层仍未标记。PV直到胎儿155 - 162天时才在第2 - 6层中出现。大星状细胞和一些锥体细胞首先出现在第5/6层和4Cα层中,但到出生后6周时,除4Cβ层外,所有层中都发现了PV+星状神经元。第4Cβ层在出生后3周时含有一些PV+细胞体,在出生后6周时神经毡染色较浅,但随后PV标记迅速增加,以至于到出生后12周时,4Cβ层的密度超过了4Cα层。到出生后20周时,纹状皮层具有成年模式的细胞数量和神经毡密度。这些发育模式表明,Cal细胞体染色的最高密度与突触形成或视觉驱动的双眼相互作用的出生后关键期无关。相反,Cal在外侧膝状体轴突到达皮层时出现,在丘脑皮质相互作用的整个过程中持续存在,并在皮层可塑性下降期间消失。PV的出现与出生时复杂视觉驱动活动的开始高度相关,而PV+细胞体的数量和PV+神经毡的密度在丘脑皮质连接完成时达到成年水平。