THE DISTRIBUTION AND PHYLOGENETIC SIGNIFICANCE OF BINUCLEATE AND TRINUCLEATE POLLEN GRAINS IN THE ANGIOSPERMS

Studies of the cytology of angiosperm pollen which extend our knowledge of the distribution of binucleate and trinucleate pollen to almost 2,000 species of flowering plants are summarized. Approximately 70% of the species studied release pollen in a binucleate stage, and none of them shed both types of pollen as a constant varietal trait. All phylogenetically primitive taxa are binucleate. The trinucleate trait evidently has originated independently at many times during angiosperm evolution. In no instance must one infer the origin of binucleate taxa from trinucleate ancestors. Most genera are monotypic with respect to pollen cytology, and only 10 genera are known to include both binucleate and trinucleate species. Among the 265 families studied, 179 include only binucleate genera and 54 include only trinucleate genera, while 32 include both types. Nearly all aquatic species with submersed flowers shed pollen in a trinucleate stage. A group of physiological differences which distinguish binucleate and trinucleate pollen is reviewed in relation to the survival and possible selective advantage of trinucleate mutants, and the significance of the binucleate pollen grain in the origin and evolution of flowering plants is discussed.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF SOUTHEASTERN NORTH AMERICA: PHILODENDRON LEAF REMAINS

Large leaves, new to the fossil record, from the Claiborne Formation in western Tennessee have been collected and analyzed. Careful analyses of venation and cuticular anatomy indicate that these fossil leaves contain specific characters found in modern species of the genus Philodendron subgenus Meconostigma. Features of venation were taxonomically more useful than other features. Reports of fossils of the Araceae are scarce and those few fossils assigned to the family frequently have been inadequately studied. This report establishes a reliable and carefully documented occurrence of the Araceae in the fossil record, and provides information concerning the differentiation of the family in time and the distributions of the genus Philodendron.     

VESSEL STRUCTURE OF GNETUM AND THE ORIGIN OF ANGIOSPERMS

Vessel structure of Gnetum gnemon L. and G. montanum Mgf. was investigated by means of light microscopy (brightfield, phase contrast and Nomarski interference optics), and scanning and transmission electron microscopy. The gyres of the first order framework are compound, consisting of three elementary strands (subunits). Pits ranging from slit-shaped to circular are formed either between the elementary strands of one gyre or between adjacent gyres. (Compound gyres are also figured for Alnus and lens-shaped pits in annular-helical elements are demonstrated for Juglans and Salix). In the two species of Gnetum studied, scalariform pits occur in addition to the well known circular pits. Scalariform perforation plates are present besides scalaroid, foraminate, and simple perforation plates as well as intermediates between the above types. These findings invalidate a common argument against the gnetalean origin of angiosperms, namely that pits and perforations of Gnetum are totally unlike those of primitive angiosperms and that therefore Gnetum must be ruled out as a potential ancestor of angiosperms. Variation in vessel structure of Gnetum is so great that it encompasses the typically circular pits of the coniferopsids as well as patterns of pits and perforation plates found in angiosperms. Some photographs of angiospermous taxa are interspersed with those of Gnetum to indicate the striking similarities. The determination of the patterns and shapes of pits and perforations is discussed in terms of four parameters: 1) the ratio of the width of the cell face to the distance between the gyres; 2) the types and distribution of the second order framework; 3) the stretching and distortion of gyres and/or second order framework; and in the case of perforations, 4) the pattern of wall and bar breakdown. Since the first parameter may change continuously, a continuum between circular pits or perforations and scalariform ones may occur. Such a continuum actually exists in Gnetum as well as in angiosperms. Patterns due to the other three of the above parameters are also formed in similar ways in both Gnetum and angiosperms. These similarities may, of course, be interpreted as the result of parallel evolution. However, when one considers the large number of angiospermous features of Gnetum, one is led to ask whether Gnetum may not have been close to the ancestral stock of all or at least some taxa of angiosperms.     

APERTURE EVOLUTION IN THE POLLEN OF PRIMITIVE ANGIOSPERMS

Diversity of pollen apertures in 35 families of the ranalean complex is compared through a series of representative scanning electronmicrographs, and the evolution of pollen aperture types in primitive angiosperms is outlined. A classification of pollen apertures found in the ranalean complex is presented, and ten basic aperture types are recognized: anasulcate, anatrichotomosulcate, zonasulculate, anaulcerate, catasulcate-cataulcerate, inaperturate, disulculate-diulculate, forate, colpate, and porate. Evidence is adduced for the primitive (ancestral) status of anasulcate pollen, and transitional stages in the evolution of other pollen aperture types in the ranalean complex are examined. From an early stock of ranalean angiosperms with anasulcate pollen, there appears to have been development of a number of interesting but evolutionally dead-end lines, represented among others by zonasulculate, anaulcerate, and catasulcate-cataulcerate pollen types. The most important evolutionary trend in early angiosperm pollen seems to have been the development of inaperturate pollen grains in many families of primitive angiosperms, from which there was a second major radiation of aperture types, including evolution of disulculate and forate pollen. Comparative study of pollen apertures observed in living primitive angiosperms suggests a de novo origin of the uniquely angiospermous (dicotyledonous) colpate pollen type from such inaperturate pollen.     

塔克拉玛干沙漠腹地3种植物根系构型及其生境适应策略

 在塔克拉玛干沙漠腹地, 采用挖掘法挖取塔克拉玛干柽柳(Tamarix taklamakanensis)、塔克拉玛干沙拐枣(Calligonum roborovskii)和罗布麻(Apocynum venetum )根系, 对根系的拓扑结构特征进行了测定与分析。结果表明: 1) 3种植物根系均以水平分布占优势, 根系浅层化。2) 3种植物根系结构的适应性不同, 表现为两种不同的根系分支模式, 塔克拉玛干柽柳根系为叉状分支结构(q_a=0.15、q_b=0.09、TI=0.658), 罗布麻(q_a=0.43、q_b=0.35、TI=0. 83)和沙拐枣(q_a=0.52、q_b=0.38、TI=0.86)根系趋向于鱼尾形分支结构。3) 3种植物根系的连接长度都较大, 最小也达1.12 m, 说明在塔克拉玛干沙漠腹地, 3种植物通过增加连接长度来扩大根系在土层中的分布范围, 从而提高根系的有效营养空间, 增加根系连接长度是根系对沙漠腹地贫瘠土壤环境的一个良好适应。4)研究验证了Leonardo da Vinci法则, 即根系分支前的横截面积等于根系分支后的横截面积之和, 3种植物根系分支前后的横截面积符合Leonardo da Vinci法则。研究表明沙漠腹地3种植物根系构型特征既有相似性又有差异性, 在相似的沙漠环境中具有不同的根系适应策略。