Distribution and Evolution of Forms and Types of Sieve-element Plastids in the Dicotyledons

Sieve-element plastids may contain any combination of protein crystals (c), protein filaments (f), and starch grains (s), or none of these. All structurally distinct combinations (=forms) possible (s, cs, cfs, c, cf, fs, f, o) are recorded among the 2100 dicotyledons investigated so far with the transmission electron microscope (representing 381 of more than 460 families described). The six forms that include c and/or f define the P-type and are found in some 620 species, mainly confined to Magnolianae, Caryophyllanae, Vitanae, and Rutanae (Fabales, Linales, Rhizophorales). A detailed family-by-family analysis and quantitative form-by-form comparison suggests that form-Ss plastids give rise to all other forms. Based on a logic model connecting all forms via one-step-alterations (loss or gain of starch, protein crystals, or filaments), the distribution of P-type plastids within dicotyledons, their interconnection, and their relationships to the S-type taxa are discussed. Due to the diversity of sieve-element plastids found within one taxon, the Magnolianae (Pand S-type) are held to be a center of P-type evolution and more primitive than Caryophyllanae (P-type, only) or other superorders. The significance of sieve-element plastid data to the shaping of some of the higher dicotyledon taxa is discussed.