Филополий

Однако конус роста нейрона С связывается всегда с поверхностью клеток Р, а не А. По-видимому, нейрон G способен не только узнать нервный пучок А/P среди 25 нервных пучков, но и отличить в пределах одного пучка один вид аксонов от другого. Именно филополий, исходящие из конуса роста нейрона С, являются теми структурами, которые первыми распознают аксоны Р. Они избирательно соединяются с поверхностью двух аксонов Р и обвиваются вокруг них. На этой стадии филополии отходят как от конусов роста, так и вбок от аксонов. К нашему удивлению, мы обнаружили, что даже в путанице переплетения многочисленных филоподнй эмбрионального нейропиля филополии нейрона С узнают и находят не только аксоны нейрона Р, но даже и его филоподии. Отсюда следует, что — независимо от того, какова природа меток, по которым нейроны распознают определенные клеточные поверхности, — эти метки расположены на всей поверхности растущего аксона.

Такой избирательный рост аксонов вдоль определенных нервных пучков, опосредуемый филоподиямн и конусами роста, играет двоякую роль в формировании нервной системы. Во- первых, эти нервные пучки, подобно магистралям, направляют конусы роста в нужное место нервной системы. Во-вторых, они создают конкретное локальное окружение для отростков каждого нейрона, среди членов которого тот впоследствии выбирает своих синаптических партнеров. На ранних стадиях развития, до того как отростки нейронов в нейропиле переплетутся настолько, что создадут непроходимый «войлок», нейроны могут контактировать и распознавать друг друга на значительных расстояниях, так как их филополии могут почти без ограничений сталкиваться друг с другом. Контакты клеточных поверхностей и создают предпосылку для избирательного движения конусов роста вдоль определенных нервных пучков. Позже, когда начинают образовываться синаптические связи, области, в пределах которых нейроны могут контактировать друг с другом, видимо, уменьшаются. Это можно объяснить тем, что увеличиваются размеры нейропиля, растет плотность составляющих его нейронных отростков и, как следствие, уменьшается пространство для движения разыскивающих свои мишени филоподий. В результате выбор нейроном потенциальных партнеров ограничивается клетками, чьи отростки проходят в нервных тяжах в непосредственном соседстве с его собственными.

 

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