Cortical structure and function in euglenoids with reference to trypanosomes, ciliates, and dinoflagellates

The membrane skeletal complex (cortex) of euglenoids generates and maintains cell form. In this review we summarize structural, biochemical, physiological, and molecular studies on the euglenoid membrane skeleton, focusing specifically on four principal components: the plasma membrane, a submembrane layer (epiplasm), cisternae of the endoplasmic reticulum, and microtubules. The data from euglenoids are compared with findings from representative organisms of three other protist groups: the trypanosomes, ciliates, and dinoflagellates. Although there are significant differences in cell form and phylogenetic affinities among these groups, there are also many similarities in the organization and possibly the function of their cortical components. For example, an epiplasmic (membrane skeletal) layer is widely used for adding strength and rigidity to the cell surface. The ER/alveolus/amphiesmal vesicle may function in calcium storage and regulation, and in mediating assembly of surface plates. GPI-linked variable surface antigens are characteristic of both ciliates and the unrelated trypanosomatids. Microtubules are ubiquitous, and cortices in trypanosomes may relay exclusively on microtubules and microtubule-associated proteins for maintaining cell form. Also, in agreement with previous suggestions, there is an apparent preservation of many cortical structures during cell duplication. In three of the four groups there is convincing evidence that part or all of the parental cortex persists during cytokinesis, thereby producing mosaics or chimeras consisting of both inherited and newly synthesized cortical components.