POLLEN PORE DEVELOPMENT AND ITS SPATIAL ORIENTATION DURING MICROSPOROGENESIS IN THE GRASS SORGHUM BICOLOR

The spatial relationships observed during microsporogenesis and pollen development in Sorghum bicolor indicate that a strong polarization exists in the anther locule and within individual microspores and pollen grains. During all developmental stages, each sporogenous cell and its derivatives lie continuously adjacent to the tapetum. The microspores and pollen grains form depressions in the tapetal orbicular wall. When the single pore of each microspore is initiated, as a gap in the primexine, it too lies adjacent to the tapetum and remains tightly appressed there until pollen maturity. A sequence of polar phenomena in microspores and pollen grains centers on an axis through the pore and perpendicular to the tapetal surface. These events include migrations of the microspore and vegetative nuclei, initial placement of the generative cell opposite the pore and its later migration, and a polar engorgement process whereby the pore end of the pollen grain (adjacent to the tapetum) fills with starch grains first. The tapetal cytoplasm completely degenerates at precisely the time of pollen engorgement, and its degradation products are believed to be available for pollen uptake at this time. The continuous association of the sporogenous cells or their cellular derivatives and their pores with the tapetum is thought to play an indispensible role in pollen development in sorghum and probably in all other grasses as well. The consistent position of the pore adjacent to the tapetum should be considered another common feature of microsporogenesis in the Gramineae. The characteristic exine pattern forms over the operculum and annulus of the pore, but the lamellae, which underlie the annulus, form a highly modified multilayered nexine. Membrane-like cores are observed in these lamellae and are believed to be involved in the initiation of sporopollenin deposition, but they are obliterated by pollen maturity. Neither the cores nor the lamellae are found in other parts of the pore or in the nonapertured wall.     

POLLEN GRAIN DEVELOPMENT AND TAPETAL CHANGES IN STRELITZIA REGINAE (STRELITZIACEAE)

The proexine that forms within the callosic envelope before the end of the microspore tetrad period is thick (about 1 μm) and exceptionally complex. It has components equatable with tectum, columellae, and a nexine that includes lamellar zones. All these components persist in the exine although late in development they become difficult to recognize because this exine is reduced in thickness, apparently by stretching, to a maximum of 0.2 μm. Strelitzia is an example of an exine template, with receptors for sporopollenin, that is not maintained during development. The Strelitzia microspore surface changes from an exine like that on an interaperture sector to the channeled intinelike system common for the apertures of pollen grains. The exine on sterile grains gives what may be a rare view of a stabilized immature exine. The mature exine on viable pollen grains resembles this early exine only in the most impressionistic way. Tapetal cells go through at least one cycle of hyperactivity, dedifferentiation, mitosis, and then again hyperactivity before they finally decline.     

POLLEN WALL AND APERTURE DEVELOPMENT IN HELIANTHUS ANNUUS (COMPOSITAE: HELIANTHEAE)

Wall development of tricolpate pollen of sunflower was studied by light and by scanning and transmission electron microscopy. The wall and colpi are initiated during the tetrad stage, producing a young, spinulate, two-layered exine (ektexine and endexine) separated by a “spacer layer.” After release from the tetrads, the individual microspores round up and enlarge. The exine layers increase in thickness and complexity from sporopollenin contributed by the tapetum and microspores. During the mid-vacuolate microspore stage, the tapetum becomes plasmodial and surrounds the developing microspores. At the vacuolate pollen stage, after the wall and colpi are completely formed, the plasmodial tapetum breaks down and releases its contents into the locule. Some of the contents are presumably utilized by the pollen to make storage reserves while other components, such as lipids and proteins, fill the spaces within the pollen wall exine. Pollen wall ontogeny provides a scheme of terms for mature composite walls in general. The various events associated with microsporogenesis in sunflower are compared with those reported in other pertinent studies.     

SPECIAL PAPER: SYSTEMATICS AND EVOLUTION OF SORGHUM SECT. SORGHUM (GRAMINEAE)

The genus Sorghum Moench is subdivided into sections Chaeotosorghum, Heterosorghum, Parasorghum, Stiposorghum and Sorghum. Section Sorghum includes two rhizomatous species, S. halepense (L.) Pers. (2n = 40) and S. propinquum (Kunth) Hitchcock (2n = 20), as well as the annual S. bicolor (L.) Moench (2n = 20). Sorghum bicolor is divided into subspecies bicolor to include all domesticated grain sorghums, subspecies drummondii (Steud.) de Wet comb. nov. to include stabilized derivatives of hybridization among grain sorghums and their closest wild relatives and subspecies arundinaceum (Desv.) de Wet et Harlan to include the wild progenitors of grain sorghums. Four ecotypes of subspecies arundinaceum are recognized: race aethiopicum of the arid African Sahel. race virgatum of northeastern Africa, race arundinaceum of the African tropical forest, and race verticilliflorum of the African Savanna. The numerous, usually recognized grain sorghums are divided among five basic races, bicolor, caudatum, durra, guinea and kafir, and ten hybrid races that each combine characteristics of at least two of these basic races. Races of grain sorghum are morphologically distinct, and they maintain their unity of type through spacial and ethnological isolation.     

MICROSPOROGENESIS AND TAPETAL DEVELOPMENT IN NORMAL AND MALE-STERILE CARROTS (DAUCUS CAROTA)

Zenkteler , Maciej . (U. Adam Mickiewicz, ul. Stalingradzka 14, Poznan, Poland.) Microsporogenesis and tapetal development in normal and male-sterile carrots (Daucus carota). Amer. Jour. Bot. 49(4): 341–348. Illus. 1962.—Meiosis and anther development proceed normally in fertile plants. Nine pairs of chromosomes are present at diakinesis and at metaphase I. The mature pollen grains possess 2 male gametes at the time of shedding; 80–92% of the pollen appears normal. A cross-shaped configuration at pachytene characteristic of a reciprocal translocation is present in the completely pollen-sterile plants indicating that one of the parents is homozygous for an interchange between 2 members of the chromosome complex. Chromosome bridges with fragments at anaphase I and anaphase II lead to aberrant chromosome distribution during meiosis. Complete microspore abortion is associated with a periplasmodium formation of the tapetum and anther wall deterioration.     

ALLOZYME VARIATION IN OLD WORLD RACES OF SORGHUM BICOLOR (POACEAE)

A survey of allozyme variation in cultivated races of sorghum (Sorghum bicolor) was undertaken. Eight plants each of 83 accessions representing the five primary races and five of the intermediate races of sorghum were analyzed for 15 enzyme systems encoded by 27 loci. Low levels of variation were found within accessions, which is typical of self-pollinating species. Little variation was also found among accessions. Compared with other cereals, S. bicolor is depauperate in allozyme variation. We found an average of 1.81 alleles per locus with a mean expected heterozygosity of 0.008 for the accessions and total panmictic heterozygosity of 0.093. Only 9% of the variation present was found within accessions, while 91% was among accessions. Most of the variation present is attributable to differences in geographic origin of the accessions rather than racial differences. Western and eastern Africa have the highest levels of total heterozygosity (0.108 and 0.088, respectively), while southern Africa has the lowest (0.008). Principal component analysis revealed continuous variation among races and geographic regions with the accessions failing to segregate into discrete racial or geographic clusters.     

ULTRASTRUCTURE AND DEVELOPMENT OF THE NEXINE AND INTINE IN THE POLLEN WALL OF SILENE ALBA (CARYOPHYLLACEAE)

Nexine and intine development in Silene alba (Caryophyllaceae) was investigated by electron microscopy and enzyme cytochemistry. Nexine-2 forms by deposition of sporopollenin along unit membrane lamellae closely associated with the microspore plasma membrane in the late tetrad stage. After the callose wall dissolves, electron density increases along the tangentially oriented fibers of the proximal primexine, forming nexine-1. When the exine is essentially complete, the intine begins to develop. In the nearly mature microspore, acid phosphatase activity appears in the peripheral cytoplasm just prior to its extrusion into the intine of the mature pollen grain.     

四倍体高粱与约翰逊草间的亲缘关系分析

本研究以四倍体高粱与约翰逊草为材料,利用SSR分子标记和细胞遗传学方法分析了高粱与约翰逊草间的亲缘关系,SSR分析结果表明,高粱与约翰逊草的遗传背景差异较大,SSR差异位点和相似位点在连锁群上的分布具不平衡性;按照差异引物出现频率高低,将连锁群分为两类：高度差异区和低度差异区。细胞学分析结果表明：（1）双亲及杂交种都是不规则的四倍体遗传群体。（2）花粉母细胞减数分裂中期I,双亲及杂交种染色体配对以二价体和四价体为主,杂交种平均每个细胞二价体数为17．00,四倍体高粱为15．23、约翰逊草为15．83,四价体数分别为0．95,2．15和1．60个。但杂交种减数分裂过程中也出现一定数量的单价体,减数分裂会形成一定比例的非整倍配子。SSR检测结果与细胞学分析结果具有一致性,约翰逊草与高粱的染色体组间存在一定程度的同源性。二者杂交不能形成稳定遗传的双二倍体。     

ANTHER AND POLLEN DEVELOPMENT IN RICE (ORYZA SATIVA)

Investigations of the growth of anthers and ontogeny of pollen grains of Oryza sativa (rice) IR-30 were undertaken for the purpose of 1) providing a set of growth measurements and 2) describing stable cytological features of anther and pollen development. Correlations exist between elongation of the floret and growth parameters of the anther such as its length, width, fresh and dry weights and cytological stage of pollen development. In the early ontogeny of the anther, hypodermal archesporial initials divide periclinally to form primary parietal cells and primary sporogenous cells. Each of the latter divides twice mitotically to generate four microspore mother cells, which undergo meiosis. The anther wall is formed by anticlinal and periclinal divisions of the primary parietal cells as well as of cells surrounding the primary sporogenous cells. Subsequent cytological features in the development of anther and pollen grains of rice have much in common with anther and pollen developmental biology of other members of Gramineae.     

PEARL MILLET (PENNISETUM AMERICANUM (L.) LEEKE) AND GRAIN SORGHUM (SORGHUM BICOLOR (L.) MOENCH) ULTRASTRUCTURE

Ultrastructural features of pearl millet (Pennisetum americanum (L.) Leeke) and grain sorghum (Sorghum bicolor (L.) Moench) caryospses were investigated with thin sections of the dry, mature grain in the transmission electron microscope, and fractured kernels in the scanning electron microscope. The pericarp of those grains is comprised of three distinct layers: epicarp, mesocarp of parenchyma cells, and endocarp of compressed cross and tube cells. Mesocarp cells of grain sorghum contain starch granules embedded in a cytoplasmic matrix. The major constituent of sorghum and millet aleurone cells are aleurone grains (protein bodies) and lipid bodies. Subaleurone cells contain a much higher proportion of protein bodies than starch granules, and the protein bodies are structurally distinct from those in the aleurone. The germ scutellar ultrastructures of the two grains were similar; protein bodies, lipid bodies, epidermal cells and parenchyma cells of the germ are described.     

POLLEN DEVELOPMENT IN OENOTHERA BIENNIS (ONAGRACEAE)

Development of the exine and viscin threads in Oenothera was studied by a combination of transmission electron microscopy and field emission scanning electron microscopy. Exine formation is initiated in the early tetrad stage by plate-like structures of preexine formed evenly around the microspore within a callosic wall. In the late tetrad stage, an endexine accumulates on white-lined lamellae underneath the preexine. After dissolution of the callosic wall, the preexine develops into beaded ektexine which is differentiated into an undulate tectum-like layer and columellae-like components. Interwoven fibrous strings connect the developing ektexine and the surface of the tapetal cells, and later develop into the viscin threads. These developmental processes imply that the columellae-like components are different in structure from the columellae of other angiosperms and that the formation of viscin threads is associated with the tapetal cells.     

金鱼草珠被绒毡层壁囊的分离与鉴定

由金鱼(Antirrhinum majus L.)草开花后的胚珠中分离出抗乙酰解、抗酶解、对强氧化剂有一定抗性但不抗二乙醇胺的细胞壁囊状结构。切片原位观察表明,它是珠被绒毡层的内壁,能被苏丹 IV 染色与金胺 O 荧光染色。它全面地包围胚囊内正在发育的胚和胚乳,仅在合点与珠孔两端开口,而在此二处分别有胚乳吸器与胚柄与外相通。这一结构的功能可能是保证营养物质定向导入胚囊以及防止珠被绒毡层分泌的水解酶侵蚀胚囊等。     

DEVELOPMENT OF WHEAT (TRITICUM AESTIVUM) POLLEN. II. HISTOCHEMICAL DIFFERENTIATION OF WALL AND UBISCH BODIES DURING DEVELOPMENT

The histochemistry of different developmental stages of the pollen wall, aperture, and Ubisch bodies of Triticum aestivum is examined with light and transmission electron microscopy. Various parts of the callosic envelope of the tetrad spores stain differentially. At the late tetrad stage, the probacules and the coat of pro-Ubisch bodies are densely stained for acidic polysaccharides, protein, and neutral polysaccharides. The protectum and the core of pro-Ubisch bodies are moderately stained. Upon release of microspores from the callosic cell envelope, the stainability for acidic polysaccharides increases in the exine and in the wall of Ubisch bodies, becoming very intense in the wall of mature pollen grains and Ubisch bodies. The stainability for neutral polysaccharides is decreased in the mature pollen wall and in the Ubisch bodies, while the stainability for protein increases. The results also indicate the probability of the presence of unsaturated lipids and the absence of free aldehydes in the pollen wall and Ubisch bodies.     

ORIGIN AND DEVELOPMENT OF POLLEN EMBRYOIDS AND POLLEN CALLUSES IN CULTURED ANTHER SEGMENTS OF HYOSCYAMUS NIGER (HENBANE)

The dedifferentiation of pollen grains of Hyoscyamus niger (henbane) into embryoids and calluses was examined by culturing identical segments of the same anther in a mineral salt-sucrose basal medium and in the basal medium supplemented with 2.0 mg/l 2,4-dichlorophenoxyacetic acid, respectively. Addition of auxin enhanced anther efficiency but did not affect the number of embryogenic pollen grains of an anther segment transformed into calluses. In anther segments cultured in the basal medium, the organogenetic part of the pollen embryoid was formed by the division of the generative cell alone, or by the division of both generative and vegetative cells. More or less similar pathways were followed by pollen grains of anther segments cultured in a medium containing auxin to form calluses. Culture of anther segments in a medium containing a high concentration of auxin (50.0 mg/l) led to a significant reduction in the yield of calluses which were formed almost entirely by the division of both generative and vegetative cells. The bearing of these observations on the role of auxin in determining the pathway of differentiation of embryogenic pollen grains in cultured anther segments is considered. The appearance of embryogenic pollen grains in close proximity to the tapetum as seen in longitudinal sections of cultured anther segments has suggested a role for a gradient of tapetal factors in embryogenic induction.     

须芒草族植物花粉形态的观察

在光学显微镜下和扫描电镜下对禾本科须芒草族（Ａｎｄｒｏｐｏｇｏｎｅａｅ）中分隶于８个亚族３４个属的３６种植物的花粉进行形态面容和比较研究。结果显示,本族植物花粉形态较为一致,花粉近形或扁球形,单萌发孔,孔,周围加厚,具盖,外壁表面散布有颗粒。这表明其是一个自然类群。总体来讲,芬烨大的演化分异,只是表面纹饰的和芬烨大小有一;定的差异。纹饰可分为三种类型粗糙型,不明显疠状突起才明显疠状突起型。花粉开矿     

POLLEN AND TAPETUM DEVELOPMENT IN DESMODIUM GLUTINOSUM AND D. ILLINOENSE (PAPILIONOIDEAE; LEGUMINOSAE)

Each of the four microsporangia has three or four wall layers, a uninucleate tapetum of various cell shapes with nuclei that remain in prophase, and 12-24 pollen mother cells (PMCs). A sterile transverse septum sometimes bisects the microsporangium. PMCs secrete callose but not uniformly, and contact among them continues through meiosis. Simultaneous cytokinesis by furrowing isolates each microspore in callose, which later disperses. The separated microspores become vacuolate, undergo mitosis to become pollen, and later become filled with food reserves. Endothecial wall thickening and tapetal dissolution occur after pollen engorgement. Calcium oxalate crystals form in tapetal cells during the sporogenous stage, reach maximum size during early meiosis, and remain prominent until tapetal dissolution.