Endosperm development: Lecture 6, 9-8-99

I. Reserve tissues: endosperm (transient and persistent), perisperm, and cotyledons.

II. Origin of the endosperm: From a second embryo? Arguments for and against.
III. Modes of endosperm development
1. nuclear - most common. Free nuclear phase followed by cellularization.
2. cellular - no free nuclear phase.
3. helobial - infrequent. Primary endosperm nucleus divides to produce two cells of
unequal size. The larger cell goes through a free nuclear phase and then cellularizes (as
in 1). The smaller cell may remain unicellular and uninucleate, or may undergo a few
mitoses to form a multinucleate cell.
IV. Genetic balance between embryo and endosperm and "imprinting" (The Endosperm Balance
Number, EBN).
1. maize ig (indeterminate gametophyte) mutant.
2. crosses in Arabidopsis to manipulate maternal:paternal genome ratios.
V. Domains of the endosperm:
Cereal endosperm - at maturity has 5 tissues/cell types
1. central starchy endosperm (CSE) - not viable at maturity.
2. sub-aleurone layer (SAL) - cells of the starchy endosperm closest to aleurone
3. aleurone layer (AL) - 1 cell thick in maize, 3 cells thick in barley. Alive at maturity and
involved in mobilization of reserves during germination.
4. basal endosperm transfer layer (BETL)- transfer of nutrients to developing endosperm.
BET1 (maize) and END1 (barley) are markers for the BETL.
5. embryo-surrounding region (ESR)- cytoplasmically dense cells around embryo. Esr1 is
expressed specifically in this region.
VI. Communication between maternal tissues and endosperm: the maize mn1 (miniature-1)
mutant.
VII. Storage reserves:
1. Endoreduplication in the endosperm
2. Carbohydrates: amylose and amylopectin (starch), mannans, and xyloglucans
3. Proteins - in vacuole or in the ER: globulins, albumins, prolamines
4. Oils
5. Phytin
VIII. Regulation of gene expression in the endosperm: maize O2 and zein genes as an example.