From a simple hermaphrodite flower, and from a complex incompatibility system unique among the flowering plants, several breeding systems have evolved in the Gramineae. Self-compatibility is the most commonplace variant and following this mutation, habitual or facultative cleistogamy is a simple evolutionary step. Separation of the sexes to different plants as in dioecism is relatively uncommon (ca. 20 genera), and gynodioecism is much less frequent still (3-4 genera). Both are seen as escape pathways from self-compatibility; the development of such pathways is discussed. Separation of the sexes to separate flowers as in monoecism is relatively common and with the variants andromonoecism and gynomonoecism is the most substantial departure from hermaphroditism in the family. These states are also interpreted as responses to self-compatibility; and though they do not generate cross-fertilization, they assist its evolution. Pathways for the evolution of these breeding systems are described. Apomixis and the breeding system best suited are discussed.I Reproductive biology in the Gramineae begins at the transition in the shoot apex from leaf production to the initiation of inflorescence primordia and the later development of floral structures. These have been well described for numerous grasses (Barnard, 1955, 1957, 1964; Bonnett, 1966; Sharman, 1960), and are mediated by photoperiod. The review of Evans (1964) elegantly reveals data on the interplay of daylength, temperature, and vernalization on inflorescence development. Floral induction and initiation may occur in the season of flowering (Evans, 1964), or in the season preceding inflorescence emergence (Mark, 1965; Hodgson, 1966). Inflorescence emergence is temperature or daylength dependent (Cooper, 1952; Connor, 1963; Heslop-Harrison, 1961). Temperature also controls anthesis, and later the release of pollen from anthers. Photoperiod, however, has other effects on the reproductive cycle. It may, for example, affect the frequency of cleistogamy in facultatively cleistogamous grasses (Langer & Wilson, 1965), or in a facultative apomict the frequency of apomictic or sexual embryo sacs (Knox, 1967; Knox & Heslop-Harrison, 1966), or depress maleness (Heslop-Harrison, 1959), or reduce the number of florets in the male inflorescence of Zea mays (Moss & Heslop-Harrison, 1968), or control protandry and protogyny (Emerson, 1924). Although these environmental influences are considerable, genetic influences may cause transient male and female sterility, or restore lost fertility in part or in whole, or promote redistribution of the sex-forms. A combination of both genetic and physiological factors may introduce difficulties into an interpretation of the breeding system of any grass. At the International Symposium on Reproduction in Flowering Plants held at Christchurch, New Zealand, in 1979, I presented a survey of the breeding systems known in the Gramineae (Connor, 1980). This present paper is complementary 1 J am grateful to colleagues and friends in several countries for helpful discussion, and especially to D. W. Clayton, Kew, T. R. Soderstrom, Smithsonian Institution, Ana M. Anton, Universidad Nacional de C6rdoba, Elizabeth Edgar and C. J. Webb, Botany Division, DSIR. 2 Botany Division, Department of Scientific and Industrial Research, Christchurch, New Zealand. ANN. MISSOURI BOT. GARD. 68: 48-74. 1981. 0026-6493/81/0048-0074/,$2.85/0 This content downloaded from 157.55.39.132 on Thu, 15 Sep 2016 05:44:04 UTC All use subject to http://about.jstor.org/terms 1981] CONNOR-REPRODUCTIVE SYSTEMS IN THE GRAMINEAE 49 to that; the bibliographies of both papers supplement each other. Here, as there, I follow the organization of tribes proposed by Hubbard (1973) for the grasses. THE FLOWER, THE SPIKELET, AND THE INFLORESCENCE The primitive grass flower is postulated as having three bracts and lodicules, six stamens, and a 1or 3-locular tristigmatic gynoecium (Schuster, 1910; Arber, 1934; Clifford, 1961). The spikelet is considered to have been many flowered, and the inflorescence is thought to have comprised many of these spikelets in a simple moreor less-branched, terminal panicle. Associated with these characteristics is the further postulate of entomophily (Schuster, 1910; Clifford, 1961; Stebbins,