DYNAMICS OF ARBUSCULE DEVELOPMENT AND DEGENERATION IN A ZEA MAYS MYCORRHIZA

A quantitative light and electron microscope study of developing and degenerating mycorrhizal arbuscules of Glomus fasciculatum in Zea mays was carried out in order to estimate three parameters during the colonization cycle. These were: 1) V_v(f,c), the fraction of the host cell volume occupied by a volume of fungus; 2) V_v(cy,c), the fraction of the host cell volume occupied by host cytoplasm; 3) S_v(pr,c), the surface-area-to-volume ratio of the host protoplast to the whole host cell. Uninfected cortical cells had an S_v(pr,c) of 0.13 μm²/μm³. As the fungus penetrates the cell wall, the protoplast invaginates, causing a decrease in protoplast volume and an increase in protoplast S_v. The S_v(pr,c) of a cell containing a mature arbuscule is 1.275 μm²/μm³. Because of the shrinkage of the protoplast, the S_v of the protoplast to its own volume rather than the original cell volume is 2.55 μm²/μm³, or almost a 20-fold increase. Total cell size is unaffected. When the arbuscule is mature, the fungus occupies 42% of the cell, with 24% as 1-μm-diam branches, and 18% as trunk. Arbuscular branch formation progresses at a linear rate and is the most important factor in causing the increased host S_v. The correlation coefficient for V_v(br,c) the volume fraction for arbuscular branches, vs. Sv(pr,c) is r = 0.932 (P < 0.001). Degeneration of the arbuscule is marked by a rapid decrease in branches, host S_v, and host cytoplasm. The trunk develops and degenerates at a slower rate than the branches.     

ADVENTITIOUS ROOT DEVELOPMENT FROM THE COLEOPTILAR NODE IN ZEA MAYS L.

Department of Botany and Bacteriology, University of Arkansas, Fayetteville, Arkansas 72701 Zea mays L. root development from the coleoptilar node was observed by light and electron microscopy. Roots developed opposite collateral vascular bundles in the coleoptilar nodal region. Three distinct histogens (stelar, cortical-protoderm, and root cap) became evident in early development. In median sections of the young roots, root cap and cortical regions formed a “hat” configuration over the stelar region. As the root matured, this “hat” developed centripetally to encapsulate the stelar region. Central core cells of the root cap were characterized by having numerous dictyosomes, amyloplasts, vacuoles, and thin cell walls. As these cells matured into outer or peripheral cap cells, the Golgi vesicles became hypertrophied. These hypertrophied vesicles contained a granular PAS-positive material which accumulated between the plasma membrane and the cell wall and formed a thick layer. As the PAS-positive material passed through the cell wall, it changed to a fibrillar texture. A PAS-positive material similar to that in the outer root cap cells was found adjacent to the outer walls of the protodermal cells. In median sections, PAS-positive material was not present in the promeristem region. Root cap cells as well as parent cortical cells were crushed as the young root forced its way through the parent tissue.     

C-BAND HETEROCHROMATIN AND DNA CONTENT IN ZEA MAYS

The number of mitotic chromosomal C-bands, the percent of the genome comprised of C-band heterochromatin, and genome size (4C DNA content) were determined for 22 North American inbred and open-pollinated lines of Zea mays. The number of C-bands ranged from 0 in Tama Knobless Flint to 18 in Zapolate Grande. The percent C-band heterochromatin ranged from 0% in Tama Knobless Flint to 16.9% in Tx601. Genome size varied over 23%: Gaspe Flint had the lowest DNA content (9.82 pg), and Zapolate Grande had the highest (12.12 pg). Genome size and the amount of heterochromatin were significantly correlated. The corn lines were assigned to five maturity zones encompassing a south-to-north range from Mexico to Canada. Significant negative correlations were detected between the amount of C-band heterochromatin and maturity zones, and between DNA content and maturity zones among the lines. It is speculated that the simultaneous selection by man for earlier maturation and plant size may be related to the lower DNA content of corn varieties adapted to higher latitudes. Such selection for larger plants may have been achieved through selection for more cells, which could result from the shorter mitotic cycle time that correlates with reduced DNA amount.     

INFLORESCENCE AND FLOWER DEVELOPMENT IN THE PIPERACEAE. II. INFLORESCENCE DEVELOPMENT OF PIPER

The inflorescence development of three species of Piper (P. aduncum, P. amalago, and P. marginatum), representing Sections Artanthe and Ottonia, was studied. The spicate inflorescences contain hundreds or even thousands of flowers, depending on the species. Each flower has a tricarpellate syncarpous gynoecium and 4 to 6 free stamens, in the species studied. No sepals or petals are present. In P. marginatum the apical meristem of the inflorescence is zonate in configuration and is unusually elongate: up to 1,170 μm high and up to 480 μm wide during the most active period of organogenesis. Toward the time of apical cessation both height and diameter gradually diminish, leaving an apical residuum which may become an attenuate spine or may be cut off by an abscission zone just below the meristem. The active apex produces bract primordia; when each is 40–55 μm high, a floral apex is initiated in its axil. Both bract and floral apex are initiated by periclinal divisions in cells of the subsurface layer. The bracts undergo differentiation rather early, while the floral apices are still developing. The last-produced bracts near the tip of the inflorescence tend to be sterile.     

ORGAN INITIATION AND THE DEVELOPMENT OF UNISEXUAL FLOWERS IN THE TASSEL AND EAR OF ZEA MAYS

The development of the unisexual male and female flowers of Zea mays from bisexual initials in both tassels and ears has been reinvestigated with SEM and TEM. The early stages of spikelet branch primordia, spikelet initiation, and early flower development are similar in both flowers, though differences in rates of growth of glumes, lemmas, and palea were detected. In both tassel and ear flowers, a pair of stamens arises opposite the lemmas and a third stamen initiates later at right angles to the first pair but from a point on the meristem below its insertion. Gynoecia develop on both tassel and ear flowers first as a ridge which overgrows the apical meristem giving rise to the stylar canal and the elongate silk. Male flowers arise in the tassel through selective vacuolation and abortion of the cells of the early gynoecium. The single female flower in each ear spikelet arises through the vacuolation and abortion of stamens in the upper flower and the repression of growth of and the eventual regression of the lower flower in each spikelet. The significance of these selective organ abortions for practical applications is discussed.     

SOME ULTRASTRUCTURAL OBSERVATIONS ON ENDODERMAL CELL DEVELOPMENT IN ZEA MAYS ROOTS

The fine structure of primary, secondary, and tertiary stages of Zea endodermal cell development was investigated. The casparian strip formed in situ in the anticlinal walls and remained at a fixed point relative to the endodermis-pericycle boundary. The only protoplasmic structure that had a constant spatial association with the developing strip was the plasmalemma. Plasmodesmata appeared to be more numerous on the tangential walls than on radial walls; only rarely were they located in the casparian strip. The suberized lamella developed on inner and outer tangential walls before it appeared on the radial walls. No cytoplasmic organelles were found to have any particular spatial association with this layer. The suberized lamella was about 0.04 μm thick except near plasmodesmata and along the adaxial margin of the casparian strip, where it was thicker. Occasionally it failed to form along the abaxial margin of the strip. The adherent affinity between plasmalemma and casparian strip was lost after the strip was covered by suberized lamella. The secondary wall became asymmetrically thickened by differential deposition of successive lamellae. A thin layer of secondary wall material extended across the floor of each pit. Pit cavities often contained mitochondria, and plasmodesmata were restricted to the pits. The plasmodesmata were constricted where they entered the thin layer of secondary wall material and where they penetrated the suberized lamella. The various stages of cell development tended to be asynchronous. No passage cells were observed. Endodermal cell development in Zea closely resembles that described for barley.     

CORTICAL ONTOGENY IN ROOTS. I. ZEA MAYS

Serial growth stages of young Zea mays primary roots were analyzed for patterns of ground meristem ontogeny. The number of cell layers in the cortex decreases from approximately 15 to 11 during early root growth. The cortex arises mostly by periclinal divisions in the outer portions of the ground meristem at levels 50–150 μm from the meristem tip, although some layers of outer cortex arise beyond 150 μm. The proendodermis contributes 3–5 cell layers to the cortex, but this contribution diminishes during early seedling growth as anticlinal divisions occur in the proendodermis. The relationship between the ground meristem and protoderm changes at the tip of the meristem during root elongation.     

THE ULTRASTRUCTURAL DEVELOPMENT OF THE DIMORPHIC PLASTIDS OF ZEA MAYS L.

The development of the dimorphic chloroplasts of Zea mays L. in adult foliage leaves is described, and a method of correlating ultrastructural stages by means of leaf chlorophyll is presented. In addition, the developmental changes in chlorophyll a/b ratio are discussed. Both the mesophyll and the bundle sheath plastids contain small grana at the earliest stages of plastid development. As the plastids enlarge, the mesophyll grana stacks increase in both length of the appressed membrane and in the number of thylakoids per granum. Initially, the grana stacks in the bundle sheath plastids also enlarge, but as the plastids approach full size, most of the membrane appression is lost. However, the remaining areas of appression in the bundle sheath plastids show an increase in the number of thylakoids in each small granum.