The extrafloral nectaries of cowpea (Vigna unguiculata (L.) Walp): I. Morphology, anatomy and fine structure.

The cowpea bears two distinctive types of extrafloral nectaries. One, on the stipels of trifoliolate leaves, consists of a loosely demarcated abaxial area (1-2 mm diameter) of widely-spaced trichomes (papillae) borne on a stomata-free epidermis, and lacking a specific vascular supply. Each trichome has up to eight apical (head) cells, two to four intermediate cells, and a single large stalk cell. The secretory faces of the apical cells bear wall ingrowths and an easily detached cuticle. The wall separating the stalk cell and the underlying epidermal cell(s) has a mean plamodesmatal frequency of 25/μm(2). The second type of nectary consists of a large elliptical mound of tissue (short and long axes about 2 mm and 4 mm) formed between a pair of flowers on an inflorescence stalk. It comprises four to eight cone-shaped subnits of secretory tissue, each with a circular secretory orifice and an individual supply of phloem, but not of xylem. Cells of the secretory tissue of the nectary subunits separate as they mature, and nectar flows to the orifice through the resulting intercellular spaces. Intact secretory cells and cellular debris are extruded into the nectar. Some of the sieve elements terminating in the inner secretory tissue exhibit open sieve pores. Each mature secretory cell contains many small (2 μm diameter) spherical protein bodies and one to three large (up to 2-3 μm diameter 15 μm long), paracrystalline bodies. These inclusions are absent or not fully developed in inner, less mature regions of the secretory tissue. Mechanisms of secretion are proposed for the two classes of nectary, including estimates of flux of sugar into the trichomes of the stipel nectary.