Ординатура / Офтальмология / Английские материалы / Retinal Development_Sernagor, Eglen, Harris, Wong_2006

astrocytes to produce VEGF. Indeed, VEGF mRNA in astrocytes increases dramatically with hypoxia (Provis, 2001). Since maturation of the vasculature reverses this process by bringing oxygen to the tissue, VEGF production is self-limiting, and astrocytic guidance of blood vessels appears to proceed as a delimited wave. Deep-layer hypoxia may also stimulate VEGF production in M¨uller cells, inducing growth of the deep vasculature (Provis et al., 1997). Glia may also guide vascular development by adding proteins to the extracellular matrix. For example, astrocytes may express fibronectin to guide retinal blood vessels (Risau and Lemmon, 1988), while astrocyte-derived cadherin limits blood vessel growth to areas containing glia (Dorrell et al., 2002).

Astrocytes may induce barrier properties in the retinal vasculature. Cultured astrocytes injected into the rat eye form aggregates that become vascularized and possess an intact blood–tissue barrier. In contrast, injected meningeal cells also aggregate and become vascularized, but do not possess a blood–tissue barrier (Janzer and Raff, 1987). Glial-conditioned medium induces barrier properties in cultured endothelial cells (Raub et al., 1992; Gardner et al., 1997). In rat, loss of astrocytes after hypoxia is correlated with a loss of barrier properties (Provis, 2001).

9.7 Concluding remarks

As this brief review of the literature has shown, interactions among glia, neurons and the vasculature, which are well documented in the adult retina, also play a significant role during retinal development. The story is incomplete, however, and many questions remain to be addressed: do radial glia in the retina guide neuronal migration? What is the definition of a mature M¨uller cell? How does neuronal activity influence glial maturation, including expression and clustering of K+ channels, expression of transporters and expression of receptors? Do retinal glia have roles in synaptogenesis? Do glial responses to retinal injury recapitulate events that occur during normal development (e.g. glial proliferation, glial trophic support of retinal neurons)? How do glial functions and properties change during aging?

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