Plants have been observed to show a range of plastic responses to environmental conditions. For example, the abundance and distribution of nutrients, as well as the presence and proximity of local competition, have been seen to result in changes in root proliferation and architecture. However, whilst some species have been witnessed displaying certain responses under given circumstances, experimental evidence suggests that responses to environmental factors can be far from simple, and sometimes counter-intuitive. Plant responses to components of the environment, and the benefit of such responses, are highly context sensitive. This thesis explores some of the real world complexities that result in the observed responses to hierarchical sets of environmental factors, and presents a theoretical model that seeks to elucidate the interplay between different factors and their effects on “optimal” behaviour by both individuals and populations. Starting with a simple one-dimensional model comprising a linearised approximation of a Gompertz growth function with nutrient patch dependent growth, the individual and combined effects of stochasticity in resource and competitor distribution are investigated. Complexity and functionality are progressively built up, with a resource dependent proliferation response, a scaling up into two-dimensions, and finally different intrinsic plant growth strategies trading growth rate against root system efficiency all introduced and investigated. Throughout the work presented in this thesis, complex and subtle behavioural responses and patterns emerge from seemingly simple models. The importance of stochasticity on individual and population level performance is also highlighted, and the results demonstrate the inability for mean-field approximations and expected results to capture the emergent behaviour.

Water is lost from higher plants via transpiration through stomatal pores the aperture of which is regulated by pairs of guard cells. Genetic engineering of the guard cell abscisic acid (ABA) signalling network that induces stomatal closure under drought stress is a key target for improving crop water use efficiency. In this study experiments were designed to investigate whether the biochemical mechanisms associated with the N-end rule pathway of targeted proteolysis could be involved in the regulation of stomatal apertures. The results indicate that the gene encoding the plant N-recognin, PRT6 (PROTEOLYSIS6), and the N-end rule pathway, are important in regulating stomatal ABA-responses in addition to their previously described roles in germination and hypoxia. Direct measurements of stomatal apertures showed that plants lacking PRT6 exhibit hypersensitive stomatal closure in response to ABA, and IR thermal imaging revealed reduced evapotranspiration under drought-stress. Together with a reduction in stomatal density, these properties result in drought tolerant plants. Plants lacking PRT6 are able to synthesis NO but their stomata do not close in response to NO suggesting that PRT6 is required for stomatal aperture responses to NO. Double mutant studies suggested that PRT6 (and by implication the N-end rule pathway) genetically interacts with known guard cell ABA signalling components OST1 and ABI1, and that it may act either downstream in the same signalling pathway or in an independent pathway. Several other enzymatic components of the plant N-end rule pathway were also shown to be involved in controlling stomatal ABA sensitivity including arginyl transferase and methionine amino peptidase activities. These results indicate that at least one of the N-end rule protein substrates which mediates ABA sensitivity has a methionine-cysteine motif at its N-terminus. A separate set of experiments were designed to investigate whether stomatal ABA-signalling pathways could have been conserved throughout land plant evolution. Cross-genetic complementation experiments were carried out to determine whether Physcomitrella stomatal apertures are able to respond to ABA and CO2 using a similar signalling pathway to that of flowering plants. The results demonstrated involvement of OST1 and ABI1 orthologues indicating that the stomata of the moss respond to ABA and CO2 using a signalling pathway that appears to be directly comparable to that of the model flowering plant Arabidopsis thaliana.

No description available.

This thesis explores the genetic control of osmotic stress response by utilising natural genetic and phenotypic variation that exists in naturally occurring Swedish populations of the common weed Arabidopsis thaliana (thale cress). The thesis describes the range of phenotypic responses to osmotic stress during very early development thereby characterising this population in terms of osmotic stress for the first time. Subsequently, regions of the genome that could be responsible for specific phenotypes are proposed where guanosine monophosphate synthetase (GMPS) is a strong candidate for future work.

No description available.

Regulation of higher plant growth and development involves the control of cell growth and division, since plant cells are immobile. A key point of plant cell cycle control is the G1 to S transition, which is promoted by CyclinD/CDK complexes. Several subgroups of D-type cyclins exist in higher plants, and the genes encoding these proteins appear to be under environmental and developmental regulation. In Arabidopsis, the CYCD3 subgroup consists of three members. The roles that these genes play in growth and development are explored, and the interaction between these genes and other factors controlling plant growth and development are investigated. A role for CYCD3;1 and its putative regulator ANT in root auxiliary meristem development is shown. However, whilst ant and cycd3;1 mutants shared some phenotypes, such as increased petal cell size, reduced leaf cell number and reduced root thickness, double mutants exhibited additive phenotypes, suggesting that there is not a strong regulation of CYCD3;1 by ANT. Supporting this, a physical interaction between ANT and a putative ANT-binding site from the CYCD3;1 promoter was not detected, and evidence of CYCD3;1 transcription regulation by ANT was weak. Supporting an alternative hypothesis, evidence of coregulation of ANT and CYCD3;1 by cytokinins in roots is provided. The expression of these genes in roots required cytokinins and appeared to be correlated. Roles for all three CYCD3s and the ERECTA (ER) kinase in the regulation of primary vascular tissue development are described, and genetic evidence of a link between CYCD3s and ER is provided. These genes appear to be required for cell division events in the procambium lineage. Furthermore, ER was also found to regulate secondary growth. Thus five novel regulators of root development have been identified, and important knowledge regarding mechanisms of lateral aerial organ size control has been gained.

No description available.

In the absence of telomerase, Saccharomyces cerevisiae telomeres erode leading to senescence. Rare cells can survive after this stage as they can elongate their telomeres utilizing homologous recombination. Two different types of survivors can be easily distinguished by Southern blot. Type I survivor cells, elongate the telomere by amplifying Y elements and require RAD51, RAD54, RAD55 and RAD57 for establishment. Type II survivors elongate their telomere by amplifying TG1-3 repeats, however, they require the following genes to be established: RAD50, MRE11 and XRS2, RAD59, SGS1 and KU80 in some cases. Both types require the gene RAD52. In this study several candidate genes were deleted individually in diploid type II survivor strains. The main aim of this work was to see if these genes were required for type II telomere maintenance. Most of these genes are not required for type II telomere maintenance at least until ~150 generations after deleting these genes. The exceptions were KU80 and RPB9. Ku80Δ strains switched to a new survivor type that is similar to type I and continued for the long term. RPB9 was required for two independent type II survivor strains to survive, whereas the third type II strain did not require this gene at ~150 generations after deleting the gene. After many generations (~ 350), this strain switched to type I. At long term propagation (~500 generations) after deletion of the candidate genes, all type II strains displayed telomere shortening until the propagation was stopped. However, Rad50Δ strains switched to type I after long term. Finally, the absence of the candidate genes did not affect the sensitivity of type II survivor strains to temperature. On the other hand, type II survivor strains with some genes deleted displayed sensitivity to UV.

Explaining the origin and maintenance of biodiversity is a central goal in ecology and evolutionary biology. Some of the most important, theoretical explanations for this diversity centre on the evolution of life histories. Comparative studies on life history evolution, have received significant attention in the zoological literature, but have lagged in plants. Recent developments, however, have emphasised the value of comparative analysis of data for many species to test existing theories of life history evolution, as well as to provide the basis for developing additional or alternative theories. The primary goal of this study was to explore existing theories of life history evolution using a dataset of demographic information in the form of matrix population models for a large number of plant species. By projecting average matrix population models for 207 plant species, life tables and fecundity schedules were obtained and, in turn, were used to estimate relevant life history parameters. These parameters were then used to explore the i) lability of life history traits in plants ii) their continuum of life history variation, iii) the evolution of senescence and iv) the significance of demographic entropy in population ecology. Elasticities and sensitivities of life history traits showed significant phylogenetic signal compared to other life history traits, although, all the values of phylogenetic signal observed were < 1 indicating that life history traits are generally labile. Eighty one percent of species in the datset had mortality curves that increased with age compared to one hundred percent of species that showed a reproductive value curve that decreases with age at the end of life. In particular, the parameters that measured pace and duration were inversely related suggesting in general, the presence of senescence in our data set. Finally, the tenets of the directionality theory based on demographic entropy were generally not confirmed. This study provides an important contribution to the life history evolution of iteroparous perennial plants and confirms existing theories on life history evolution.

Chez les eucaryotes, les gènes d’ARNr 45S sont présents en un grand nombre de copies organisées dans des régions chromosomiques appelées NOR pour « Nucleolus Organizer Region ». Cependant, seule une fraction de ces gènes est activement exprimée et leur activation/répression est majoritairement contrôlée par des mécanismes épigénétiques. Parmi les facteurs requis pour l’expression de ces gènes, se trouve la nucléoline, une protéine majeure du nucléole. Chez A. thaliana, la protéine NUC1 est nécessaire pour le maintien de la méthylation et le control de l’expression de variants spécifiques des gènes d’ARNr. De manière intéressante, contrairement aux animaux et aux levures, le génome des plantes possède un deuxième gène codant la nucléoline : NUC2. Au cours de cette étude, nous avons montré que les deux gènes NUC1 et NUC2 sont nécessaires pour la survie de la plante. L’étude de plantes mutées pour le gène NUC2 a révélé que cette protéine est impliquée dans l’organisation et l’expression des ADNr mais par des mécanismes antagonistes à ceux de son homologue NUC1. En effet, l’absence de la NUC2 induit une hyperméthylation des ADNr ainsi qu’une réorganisation spatiale et une variation du nombre de copie des différents variants des gènes d’ARNr. Par ailleurs, la protéine NUC1 se lie aux gènes actifs alors que la protéine NUC2 est associée à la chromatine condensée en périphérie du nucléole. En parallèle, nous avons montré que l’expression des ADNr est affectée en réponse à la chaleur et que le gène NUC2 est fortement induit. L’ensemble de ces données suggèrent un potentiel rôle de la NUC2 dans la répression des gènes d’ARNr au cours du développement et en réponse au stress. / In eukaryotes, 45S rRNA genes are highly repeated and localize in chromosomal regions known as NOR for “Nucleolus Organizer Regions”. However, only a small proportion of these genes is transcriptionally active and their activation and/or repression depends on epigenetic mechanisms. One of the factors involved in rDNA expression is nucleolin, a major nucleolar protein. In A. thaliana, nucleolin protein NUC1 is required to maintain rDNA methylation and control expression of specific rDNA variants. Interestingly, in contrast to animals and yeast, plants encode a second nucleolin gene: NUC2. Here, we show that NUC1 and NUC2 genes are both required for plant survival. Analysis of nuc2 mutant plants reveals that NUC2 protein is required for rDNA organization and expression but with mechanisms antagonistic to those described for its homologue NUC1. In fact, loss of NUC2 induces rDNA hypermethylation and a spatial reorganization of rRNA genes with changes in copy numbers of rDNA variants. Moreover, NUC1 protein binds transcriptionally active rRNA genes while NUC2 protein associates with condensed chromatin in the periphery of the nucleolus. Furthermore, we show that rRNA gene expression is affected in response to heat shock and that the NUC2 gene is strongly induced. Altogether, our results suggest a potential role of NUC2 protein in rDNA repression during development and/or in response to stress.