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.     

DISTRIBUTION OF TARTARIC ACID IN THE LEAVES OF CERTAIN ANGIOSPERMS

Stafford , Helen A. (Reed Coll., Portland, Oregon.) Distribution of tartaric acid in the leaves of certain angiosperms. Amer. Jour. Bot. 46(5): 347–352. 1959.—An optically active isomer of tartaric acid has been definitely identified and quantitatively analyzed in the leaves of 9 species of angiosperms, and trace amounts have been tentatively identified in about 22 others out of 49 species examined. In species where identification was positive, the quantity of tartrate varied from 5 to 200 μmoles/g. fresh wt. of leaf tissue; identification was based on paper chromatography and on the metavanadate colorimetric test. The 9 species include Vitis vinifera, V. labruscana, V. californica, Parthenocissus tricuspidata, P. quinquefolio, Pelargonium hortorum, Bauhinia malabarica, Phaseolus vulgaris, and Coleus blumei. Optical rotation data of the isolated acid indicate that the tartrate in Pelargonium and Parthenocissus quinquefolio is the (+) -isomer similar to that reported for Vitis vinifera, but the opposite to that reported for Bauhinia reticulata. The relationship of tartrate to other organic acids is discussed.