GAMETOPHYTES AND EMBRYOS OF ACTINOSTACHYS PENNULA,A. WAGNERI,AND SCHIZAEA ELEGANS,WITH NOTES ON OTHER SPECIES

Gametophytes and embryos of Actinostachys wagneri, A. pennula, and Schizaea elegans are described along with a few observations on those of 5. dichotoma, A. digitata, A. spirophylla, Ophioglossum pendulum, Danaea simplicifolia, Botrychium virginianum, B. dissectum, and Psilotum nudum. Variation in the gametophyte and embryo at the interspecific level within the Schizaeaceae is discussed.     

LEAF DEVELOPMENT OF PHASEOLUS VULGARIS L. IN LIGHT AND IN DARKNESS

Development of the primary bean leaf in the dark and under continuous white light was studied during 14 days after sowing. The increase in surface area of the blade is the result of a number of sequential processes. Both in the darkness and under illumination, leaf growth is characterized by an initial cell enlargement followed by intensive cell division. Cell division in etiolated leaves continues for one day longer than in illuminated ones, but it proceeds at a slower rate. Mature leaves grown under white light undergo a phase of cell enlargement after cell division has stopped. This increases their surface area up to 800 times when compared with the blade area of the embryo. This enlargement phase is almost absent in dark-grown seedlings. Consequently the blade area of etiolated leaves is only 50 times that of the embryonic state. Thus light appears to have a dual effect on leaf development: it activates cell division and induces cell expansion.     

GROWTH AND GRAVITATIONAL RESPONSE IN THE DEVELOPMENT OF LEAF BLADE HYPONASTY

Rapid (4 hr) auxin-induced hyponastic curvature of primary leaves of Phaseolus vulgaris is shown to depend on a positive increase in growth of the lower portion of the blade. The curvature involves laminar growth as well as vein growth and is not due to simple turgor changes. The response is sensitive to gravitational orientation, as inversion and horizontal rotation reduce the auxin-induced curvature. The ethylene-generating compound, 2-chloroethylphosphonic acid, had no hyponastic effect on the leaves when applied to either the upper or lower surface and it inhibited auxin-induced hyponasty. This inhibition was additive to that of inversion. Long-term (24–48 hr) effects of 1 mM auxin depend on the surface of the leaf treated. Application to the upper surface results in epinasty, lower surface application in hyponasty, although the initial response in each case is a hyponastic curvature. A dorsi-ventral auxin transport system and differential auxin sensitivity of upper and lower portions of the leaf blade are postulated to account for these responses.     

CYTOKININ-GIBBERELLIN REGULATION OF SHOOT DEVELOPMENT AND LEAF FORM IN TOBACCO PLANTLETS

Callus and plantlets derived from callus cultures of Nicotiana tabacum var. Wisc. #38 were grown on medium containing serial combinations of gibberellic acid (GA₃) and the cytokinin 6-(3-methyl-2-butenylamino)purine (2iP). Increasing levels of both growth substances resulted in the production of greater amounts of both callus and shoot tissue. More buds were induced when the cytokinin level was increased, and this effect was counteracted by raising the GA₃/2iP ratio. Furthermore, the size and form of the shoots depended on the GA₃/2iP ratio. High ratios resulted in tall, spindly plants with narrow leaves while low ratios resulted in short shoots with rounded leaves.     

SPORE GERMINATION AND EARLY DEVELOPMENT OF THE GAMETOPHYTE OF SCHIZAEA PUSILLA

During germination of the spore of Schizaea pusilla, the first division of the protoplast was perpendicular to the polar axis and resulted in the formation of the rhizoid. The next division parallel to the polar axis of the spore gave rise to the protonemal initial. Following this “Vittaria”-type germination, the protonema that developed was characterized by an extensive branching to produce uniseriate filaments and rhizoidophores.     

芦荟叶内芦荟素细胞的发育和蒽醌类物质的积累

应用石蜡切片、半薄切片、组织化学和荧光显微镜观察相结合的方法研究了木立芦荟叶内芦荟素细胞的发生、发育以及其蒽醌类物质的积累过程。结果表明,在叶内原形成层束分化成维管束初期,原形成层束外侧的一层细胞发育成维管束鞘。原生韧皮部筛管产生时,其外方尚保留1-2层原形成层细胞,当后生韧皮部和木质部开始分化时,此层细胞分裂。在后生韧皮部和木质部发育成熟过程中,这些细胞体积逐渐增大,并液泡化,发育成为大型薄壁细胞(芦荟素细胞),位于筛管外侧。据此,芦荟叶维管束内的大型薄壁细胞的来源与韧皮部相同,属于特化的韧皮部薄壁组织细胞。用醋酸铅处理过的上述材料的切片观察表明,芦荟素细胞在细胞体积增大,并液泡化时,在液泡内出现蒽醌类物质沉淀物,在成熟细胞的大液泡中充满沉淀物,此时,在荧光显微镜下芦荟素细胞发出桔黄色荧光。可见,此种芦荟素细胞是芦荟叶内蒽醌类物质的主要储存场所。     

STRUCTURE AND DEVELOPMENT OF SIEVE ELEMENTS IN THE LEAF OF ISOETES MURICATA

Leaf tissue of Isoetes muricata Dur. was fixed in glutaraldehyde and postfixed in osmium tetroxide for electron microscopy. The very young sieve elements can be distinguished from contiguous parenchyma cells by their distinctive plastids and the presence of crystalline and fibrillar proteinaceous material in dilated cisternae of the rough ER. During differentiation, the portions of ER enclosing this proteinaceous substance become smooth surfaced and migrate to the cell wall. Along the way they apparently form multivesicular bodies which then fuse with the plasmalemma, discharging their contents to the outside. At maturity, the sieve element contains an elongate nucleus, which consists of dense chromatin material, and remnants of the nuclear envelope. In addition, the mature sieve element is lined by a plasmalemma and a parietal, anastomosing network of smooth ER. Both plastids and mitochondria are present. P-protein is lacking at all stages of development. Tonoplasts are. not discernible in mature sieve elements. The end walls of mature sieve elements contain either plasmodesmata or sieve pores or both, but only plasmodesmata occur in the lateral walls.     

LEAF DEVELOPMENT,PHOTOSYNTHESIS, AND C14 DISTRIBUTION IN POPULUS DELTOIDES SEEDLINGS

Rates of net photosynthesis and dark respiration and distribution of C¹⁴ from selected leaves were determined for young cottonwood (Populus deltoides) trees at different stages of development. Four series of five trees—one series for each of four treated leaf positions—were included in the study. Maximum C¹⁴ export occurred when a leaf had just attained maximum size. Lower stem leaves reached maturity quickly and began exporting photosynthate when demands of the young seedling were high. Leaves at higher stem positions matured more slowly, but senescence was also delayed so their effective export life was prolonged. Translocation from a newly exporting leaf was primarily upward to developing leaves and the apex. As a leaf at any one position aged, the translocation pattern gradually shifted from upward to bi-directional and finally to a predominantly downward direction. Photosynthate translocated downward was incorporated into stem wood and roots. Maximum photosynthetic efficiency coincided with the downward shift of C¹⁴ export. Thereafter, net photosynthesis began to decline, at first slowly and then more rapidly. The patterns of photosynthesis, respiration, and C¹⁴ export associated with leaf age all varied according to leaf position on the stem.     

LEAF DEVELOPMENT IN ISOPHYLLOUS AND FACULTATIVELY ANISOPHYLLOUS SPECIES OF PENTADENIA (GESNERIACEAE)

Shoots of Pentadenia orientandina exhibit varying degrees of anisophylly, ranging from pairs of equal-sized leaves to pairs of large ventral and small dorsal leaves. In this study we compare phyllotaxis, leaf expansion, and accompanying histological changes in extremely anisophyllous shoots of this species and in isophyllous shoots of the related species, P. crassicaulis. In P. orientandina, decussate phyllotaxis is modified at leaf initiation, and angles of leaf insertion appear to be further changed during leaf expansion. In both species, leaf primordia of a pair are not distinguishable at inception, suggesting an equivalent developmental potential. In P. orientandina, size differences between ventral and dorsal leaves become significant at the P2 or P3 stage, coincident with lamina initiation. Minute dorsal leaves are arrested in their development at the P3 stage and mature without differentiation of multiple epidermis, stomata, mesophyll and most vascular tissue. Variation in dorsal leaf structure in P. orientandina emphasizes the plasticity of leaf development in this facultatively anisophyllous species.     

A GENETIC SYNDROME AFFECTING LEAF DEVELOPMENT IN PISUM

A recessive foliage mutant of Pisum, designated ‘sinuate leaf’ (sil), was found to have two distinct forms of expression, depending on the background genotype. In an af/af background—wherein leaflets are converted to tendrils—sil/sil plants had adventitious tendrils arising from clefts in the distal portion of the stipule. These adventitious tendrils were morphologically modified, just as were the true tendrils on the same plant, by different allelic combinations at the tl locus. In the standard Af background, sil/sil plants had neither incised stipules nor adventitious tendrils, although they did have undulated and somewhat distorted leaflets and stipules. Because mutant expression was variable in an Af background, classification of segregating populations was uncertain. This uncertainty was removed by taking advantange of pleiotropic effects exerted by sil in the presence of one of the wax mutants, wlo, wb, or wsp. Homozygous wlo plants ordinarily have uniformly waxy stipule surfaces, but when the plants were also homozygous for sil the stipule tips were waxless. Conversely, wb/wb and wsp/wsp plants ordinarily have uniformly waxless stipules, but when wb/wb or wsp/wsp plants were also homozygous recessive for sil the stipule tips were waxy. However, sil had no observable effects of any kind on the stipule tips of plants with stipules reduced in size by the action of st/st. By their individual and combined effects, the foliage mutants used in this study revealed developmental relationships among leaf parts not otherwise evident in non-mutant plants.     

BIOCHEMICAL AND PHYSIOLOGICAL ASPECTS OF LEAF DEVELOPMENT IN COCOA (THEOBROMA CACAO)

The dormant phase of the flush cycle of leaf growth in cocoa is known to be correlated with high abscisic acid (ABA) levels in the mature leaves of the new flush (NF) and previous flush (PF) leaves. Defoliation of either the NF leaves or PF leaves of cocoa seedlings reduced the length of the dormant phase of the flush cycle, thus showing that the mature leaves were a source of growth inhibitors which could affect shoot apical activity. The application of ABA to the NF and PF leaves led to an extension of the dormant phase, whereas application of zeatin or gibberellic acid decreased it. The distribution of [¹⁴C]ABA following its application to NF and PF leaves at different stages throughout the growth cycle showed that [¹⁴C] ABA was accumulated by the bud in relatively larger amounts during the final stages of bud dormancy (I-1 and I-2) than in the earlier stage (F-2). The results suggest that internal competition for nutrients may be responsible for the inhibition of growth at the F-2 stage but that ABA translocated from the mature leaves causes the buds to remain dormant during the subsequent stages of I-1 and I-2.     

LEAF DEVELOPMENT IN DOXANTHA UNGUIS-CATI (BIGNONIACEAE)

Leaf structure in Doxantha unguis-cati is polymorphic. The usual mature compound leaf is composed of two lanceolate leaflets and a terminal tripartite spine-tendril. Leaf primordia are initiated simultaneously in pairs on opposite flanks of the shoot apical meristem by periclinal cell divisions in the third subsurface layer of the peripheral flank meristem. Two leaflet primordia are the first lateral appendages of the compound leaf. Initiation of these leaflet primordia occurs on the adaxial side of a compound leaf primordium 63–70 μm long. Lamina formation is initiated at the base of a leaflet primordium 70–90 μm long and continues acropetally. Mesophyll differentiation occurs in later stages of development of leaflets. The second pair of lateral appendages of the leaf primordium differentiate as prongs of the tendril. Initiation of the second pair of lateral appendages occurs on the adaxial side of a primordium approximately 168 μm long. Acropetal procambialization and vacuolation of cells extend to the apex of tendrils about 112 μm long, restricting the tendril meristem to the adaxial side of the primordium and resulting in curvature of the tendril. The tendril meristem is gradually limited to a more basipetal position as elongation of apical cells continues. Initiatory divisions and early ontogenetic stages of leaflets and tendrils are similar. Their ontogeny differs when the lateral primordia are approximately 70 μm long. Marginal and submarginal initials differentiate within leaflets but not in tendrils. Apical growth of tendrils ceases very early in ontogeny as compared with leaflets.     

THE INFLUENCE OF WATER STRESS AND TEMPERATURE ON LEAF PUBESCENCE DEVELOPMENT IN ENCELIA FARINOSA

The degree of leaf pubescence development in the arid land shrub Encelia farinosa Gray is affected by air temperature, leaf water potential, and previous history of the apical meristem during the current growing season. Changes in leaf pubescence levels change leaf spectral characteristics and affect both leaf temperature and photosynthesis. Decreasing leaf water potentials and increasing air temperatures both independently increase pubescence development as measured by decreased leaf absorptances. During any one growing season leaf absorptance may change reversibly coincident with air temperature changes, but with respect to water stress leaf absorptance only decreases as the season progresses. The ecological significance of regulation of the leaf spectral characteristics is discussed.     

HETEROCHRONY AND HETEROBLASTIC LEAF DEVELOPMENT IN TWO SUBSPECIES OF CUCURBITA ARGYROSPERMA (CUCURBITACEAE)

To date, reports of paedomorphosis at the whole plant or shoot level have been loosely based on whole plant form or on the sequence of leaf shapes produced along the shoot (heteroblasty). However, interpreting the significance of heterochrony in the evolutionary loss or gain of heteroblasty based on mature leaf forms assumes that all leaves with the same shape arose through very similar modes of organogenesis. This study examines this assumption in two subspecies of Cucurbita argyrosperma, one that is wild and heteroblastic and a second that is cultivated and not markedly heteroblastic. All leaves of the cultivar are visually similar to early leaves of the wild subspecies. The cultivar is considered to be the progenitor of the wild subspecies. Scanning electron microscopy and allometry of developing leaves showed that at early nodal positions along the primary shoot, leaf development in both subspecies was similar. At later nodal positions, very young leaves of both subspecies were more similar to each other than to leaves at earlier nodal positions within the same plant at the same stage of development. This early similarity was masked in the mature shapes of later leaves due to subsequent differences in allometric growth. Thus a simple hypothesis of paedomorphosis in which the early leaf form in the progenitor is simply reiterated at later nodal positions in the cultivar is not supported by patterns of leaf development.