Cell-cell interactions during plant development.
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The crucifer Arabidopsis thaliana has been used widely as a model organism for the study of plant development. We describe here the development of an efficient insertional mutagenesis system in Arabidopsis that permits identification of genes by their patterns of expression during development. Transposable elements of the Ac/Ds system carrying the GUS reporter gene have been designed to act as enhancer traps or gene traps. A novel selection scheme maximizes recovery of unlinked transposition events. In this study 491 plants carrying independent transposon insertions were generated and screened for expression patterns. One-half of the enhancer trap insertions and one-quarter of the gene trap insertions displayed GUS expression in seedlings or flowers, including expression patterns specific to organs, tissues, cell types, or developmental stages. The patterns identify genes that act during organogenesis, pattern formation, or cell differentiation. Transposon insertion lines with specific GUS expression patterns provide valuable markers for studies of Arabidopsis development and identify new cell types or subtypes in plants. The diversity of gene expression patterns generated suggests that the identification and cloning of Arabidopsis genes expressed in any developmental process is feasible using this system.
Recent research has demonstrated that DNA methylation plays an integral role in regulating the timing of flowering and in endosperm development. The identification of key genes controlling these processes, the expression of which is altered in plants with low methylation, opens the way to understanding how DNA methylation regulates plant development.
The canavanine content of developing leaves of jack bean, Canavalia ensiformis (L.) DC., increases during leaf development. The leaf possesses the enzymes required for synthesizing canavanine by a cyclic series of reactions analogous to the ornithine-urea cycle. This reaction series involves the sequential formation of canaline, O-ureidohomoserine, and canavaninosuccinic acid.
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The combined expression of the rol A, B, and C loci of Agrobacterium rhizogenes Ri-plasmids establishes, in transgenic tobacco plants, a pathological state called hairy-root syndrome. However, when expressed separately they provoke distinct developmental abnormalities characteristic for each rol gene. Moreover, changes in their mode of expression obtained by replacing the promoters of the rol B and C genes with the cauliflower mosaic virus 35S promoter elicit new and distinct developmental patterns. These results indicate that the different rol gene products have either different targets, or have a qualitatively different effect on the same target. The target(s) must be involved in the control of plant development. Although each of the three rol genes are independently able to promote root formation in tobacco, efficient root initiation and growth is best achieved through the combined activities of more than a single rol gene. Models explaining the biological effects of A. rhizogenes-derived TL-DNA genes are discussed.
A key problem in developmental biology is understanding the origin of morphological innovations. Comparative studies in plants with different leaf morphologies indicate that the developmental pathway defined by KNOTTED1-type homeodomain proteins could be involved in generating different leaf forms. The differential expression of regulatory proteins has emerged as an important factor in driving morphological innovations in the plant kingdom--an idea that is well supported by quantitative trait locus analyses.
knox genes encode homeodomain-containing transcription factors that are required for meristem maintenance and proper patterning of organ initiation. In plants with simple leaves, knox genes are expressed exclusively in the meristem and stem, but in dissected leaves, they are also expressed in leaf primordia, suggesting that they may play a role in the diversity of leaf form. This hypothesis is supported by the intriguing phenotypes found in gain-of-function mutations where knox gene misexpression affects leaf and petal shape. Similar phenotypes are also found in recessive mutations of genes that function to negatively regulate knox genes. KNOX proteins function as heterodimers with other homeodomains in the TALE superclass. The gibberellin and lignin biosynthetic pathways are known to be negatively regulated by KNOX proteins, which results in indeterminate cell fates.
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Polycyclic aromatic hydrocarbons (PAHs), ubiquitous environmental pollutants, are known to be biologically active in mammalian systems and are accumulated by plants. A few reports suggest that PAHs stimulate growth in plants and induce morphogenesis in plant tissue in culture. To investigate the growth altering abilities of PAHs in plants, polypodiaceous fern gametophytes were grown under sterile conditions on media containing a biologically active PAH, benzo[a]pyrene (BaP), at doses ranging from 0.1 to 10.0 micrograms/ml. The growth pattern of polypodiaceous fern gametophytes enables alterations in growth and morphogenesis to be observed at the cellular level in an intact plant. Doses of BaP in the range 0.1--3.2 micrograms/ml enhanced the onset of the morphological transitions from 1-dimensional (1D) to 2-dimensional (2D) growth. This transition for BaP treated plants occurred after fewer cell divisions than the corresponding solvent and untreated controls. The low (0.1 and 0.32 micrograms) and high (1.0 and 3.2 micrograms) doses of BaP were found to accelerate and inhibit cell proliferation, respectively. The 10.0 microgram dose was toxic and resulted in decreased germination of spores and reduced survival of plants. A slight but significant decrease in survival was also observed in ferns treated with 3.2 micrograms. This is the first example of PAH influencing cell differentiation in a whole plant system.
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