The number of angiosperm species for which nuclear DNA amount estimates have been made has nearly trebled since the last collected lists of such values were published, and therefore, publication of a more comprehensive list is overdue. This paper lists absolute nuclear DNA amounts for 753 angiosperm species. The data were assembled primarily for reference purposes, and so the species are listed in alphabetical order, as this was felt to be more helpful to cyto- and biochemists whom, it is anticipated, will be among its major users. The paper also reviews aspects of the history, nomenclature, methods, accuracy and problems of nuclear DNA estimation in angiosperms. No attempt is made to reconsider those aspects of nuclear DNA estimation which have been fully revised previously, although the bibliography of such aspects is given. Instead, the paper is intended as a source of basic information regarding the terminology, practice and limitations of nuclear DNA estimation, especially by Feulgen microdensitometry, as currently practiced.

Nuclear DNA Amounts in Angiosperms

Many components of cell and nuclear size and mass are correlated with nuclear DNA content in plants, as also are the durations and rates of such developmental processes as mitosis and meiosis. It is suggested that the multiple effects of the mass of nuclear DNA which affect all cells and apply throughout the life of the plant can together determine the minimum generation time for each species. The durations of mitosis and of meiosis are both positively correlated with nuclear DNA content and, therefore, species with a short minimum generation time might be expected to have a shorter mean cell cycle time and mean meiotic duration, and a lower mean nuclear DNA content, than species with a long mean minimum generation time. In tests of this hypothesis, using data collated from the literature, it is shown that the mean cell cycle time and the mean meiotic duration in annual species is significantly shorter than in perennial species. Furthermore, the mean nuclear DNA content of annual species is significantly lower than for perennial species both in dicotyledons and monocotyledons. Ephemeral species have a significantly lower mean nuclear DNA content than annual species. Among perennial monocotyledons the mean nuclear DNA content of species which can complete a life cycle within one year (facultative perennials) is significantly lower than the mean nuclear DNA content of those which cannot (obligate perennials). However, the mean nuclear DNA content of facultative perennials does not differ significantly from the mean for annual species. It is suggested that the effects of nuclear DNA content on the duration of developmental processes are most obvious during its determinant stages, and that the largest effects of nuclear DNA mass are expressed at times when development is slowest, for instance, during meiosis or at low temperature. It has been suggested that DNA influences development in two ways, directly through its informational content, and indirectly by the physical-mechanical effects of its mass. The term 'nucleotype' is used to describe those conditions of the nucleus which effect the phenotype independently of the informational content of the DNA. It is suggested that cell cycle time, meiotic duration, and minimum generation time are determined by the nucleotype. In addition, it may be that satellite DNA is significant in its nucleotypic effects on developmental processes.

Nuclear DNA Content and Minimum Generation Time in Herbaceous Plants

Summary
The gross form of the nuclear genome varies greatly among plant species in both anatomy and genetic organization. Chromosome number (ft) ranges from 2 to over 600, and ploidy from 1 to over 20. The amount of DNA in the unreplicated haplophase genome (the 1C value) differs by more than 2500-fold among angiosperms. Although it has been questioned since the 1930s whether such variation is of adaptive significance and whether it is related, perhaps causally, with environmental factors, no direct or causal links have yet been found. However, variation in DNA C-value has far-reaching biological consequences and can be of considerable adaptive and hence ecological significance. Strikingly precise interspecific relationships exist between DNA C-value and many diverse phenotypic characters at the cellular level, and DNA can affect the phenotype in two ways, firstly by expression of its genie content and, secondly, by the biophysical effects of its mass and volume, the latter defined as nucleotypic effects. Nucleotypic variation in DNA C-value sets absolute limits to both the minimum size and mass of the basic unit of plant anatomy (i.e. the cell) and the minumum time needed to produce a similar cell with newly synthesized organic molecules. Moreover, in complex multicellular vascular plants, such effects at successive cell cycles are additive, so that DNA C-value influences many characters, including growth rate, seed weight, minimum generation time and type of life-cycle. Thus, the nucleotype profoundly affects where, when and how plants grow. Selection for a particular genomic form acting on its spatial or temporal consequences may occur at various levels ranging from the cell to the whole organism and may operate throughout the life-cycle or at just one stage. DNA C-value is often indirectly related to environmental factors which determine time-limited environments via selection acting on the temporal phenotypic consequences of nucleotypic variation. However, in the case of radio-sensitivity, selection for a low DNA C-value may act directly on the nucleotype itself, as the size of the nuclear DNA target directly affects the ability of the plant to survive.

Variation in genomic form in plants and its ecological implications

A survey of work on meiotic duration in diploid plants shows that the duration is positively correlated with the DNA content per nucleus and with the mitotic cycle time. However, meiotic duration is not solely determined by the DNA content per nucleus but is also affected by chromosomal organization, DNA structure and the developmental pattern of the organism. Thus, in three polyploid plant species meiosis is much shorter and in three animal species it is much longer than would be expected in diploid plant species having corresponding DNA contents. Differences in meiotic duration in plant species are usually the result of proportional differences in all the stages of meiosis. Factors affecting the initiation, control and duration of meiosis are discussed. The consequences of changes in nuclear DNA content on developmental processes and the life cycle in plants are considered. It is suggested that DNA influences development in two ways, first directly through its informational content, and second indirectly by the physical mechanical effects of its mass independent of its informational content.

The duration of meiosis

In the genomes of chromosomal organisms, cytological evidence from disparate sources suggests that each unit of information encoded as a DNA base sequence is serially repeated. Further cytological and genetical evidence suggests that among such serially repeated sequences a terminal unit serves as the ‘master’ sequence, within which recombinational events can occur, followed by ‘slave’ sequences which are not directly involved in recombination but which are made congruent to the master sequence once per life-cycle. The formation of lateral ‘lamp-brush’ loops in meiotic prophase, after synapsis, is claimed to represent the outcome of the master/slave matching process.

J. McLeish

Papers

Measurements of deoxyribosenucleic acid (DNA) in higher plants by Feulgen photometry and chemical methods

Nucleic acid content and concentration in root cells of higher plants.

The quantitative cytochemical estimation of arginine.