Epigenetic Modifications in Plant Development and Reproduction

Plants are exposed to highly fluctuating effects of light, temperature, weather conditions, and many other environmental factors throughout their life. As sessile organisms, unlike animals, they are unable to escape, hide, or even change their position. Therefore, the growth and development of plants are largely determined by interaction with the external environment. The success of this interaction depends on the ability of the phenotype plasticity, which is largely determined by epigenetic regulation. In addition to how environmental factors can change the patterns of genes expression, epigenetic regulation determines how genetic expression changes during the differentiation of one cell type into another and how patterns of gene expression are passed from one cell to its descendants. Thus, one genome can generate many ‘epigenomes’. Epigenetic modifications acquire special significance during the formation of gametes and plant reproduction when epigenetic marks are eliminated during meiosis and early embryogenesis and later reappear. However, during asexual plant reproduction, when meiosis is absent or suspended, epigenetic modifications that have arisen in the parental sporophyte can be transmitted to the next clonal generation practically unchanged. In plants that reproduce sexually and asexually, epigenetic variability has different adaptive significance. In asexuals, epigenetic regulation is of particular importance for imparting plasticity to the phenotype when, apart from mutations, the genotype remains unchanged for many generations of individuals. Of particular interest is the question of the possibility of transferring acquired epigenetic memory to future generations and its potential role for natural selection and evolution. All these issues will be discussed to some extent in this review.

Epigenetic Modifications in Plant Development and Reproduction

Plants are exposed to highly fluctuating effects of light, temperature, weather conditions and many other environmental factors throughout their life. As sessile or-ganisms, unlike animals, they are unable to escape, hide or even change their position. Therefore, the growth and development of plants is largely determined by interaction with the external environment, the success of this interaction depends on the ability of the phenotype plasticity, which is largely determined by epigenetic regulation. In addi-tion to how environmental factors can change the patterns of genes expression, epige-netic regulation determines how genetic expression changes during the differentiation of one cell type into another, and how patterns of gene expression are passed from one cell to its descendants. Thus, one genome can generate many 'epigenomes'. Epigenetic modifications acquire special significance during the formation of gametes and plant reproduction, when epigenetic marks are eliminated during meiosis and early embry-ogenesis and later reappear. However, during asexual plant reproduction, when meio-sis is absent or suspended, epigenetic modifications that have arisen in the parental sporophyte can be transmitted to the next clonal generation practically unchanged. In plants that reproduce sexually and asexually, epigenetic variability has different adap-tive significance. In asexuals, epigenetic regulation is of particular importance for im-parting plasticity to the phenotype, when the genotype remains unchanged for many generations of individuals. Of particular interest is the question of the possibility of transferring acquired epigenetic memory to future generations and its potential role for natural selection and evolution. All these issues will be discussed to some extent in this review. In the last two decades, a lot of data on the epigenetic regulation of plants has appeared, as well as works summarizing the accumulated knowledge (Verhoeven and Preite 2013; Pikaard and Scheid 2014; Gehring 2019; Ono and Kinoshita 2021), nevertheless, many questions remain unclear, and a number of results are contradic-tory. New in this area data is constantly emerging. We tried to take into account and discuss the main findings and conclusions in this field.

Expected response to early-generation selection for yield and tuber appearance traits in potatoes

The aim of this study was to evaluate the performance of potato clonal families, and to estimate genetic variance, heritability and the expected response to selection of tuber yield and appearance traits in early generations. Twelve potato families were obtained from crosses between two groups of randomly-selected genotypes, including Eliza, C1730-7-94, and C-1742-8-95 in group 1; and Shepody, Asterix, Caesar, and White Lady in group 2. The crosses were made in factorial design (3 genotypes x4 genotypes), and each family consisted of 75 genotypes. Experiments were conducted in the fall of 2010, with a seedling generation under greenhouse conditions, and in the fall of 2011, with a clonal generation under field condition. High heritability estimates suggest that mild to moderate selection can be applied in the seedling generation to eye depth, eyebrow prominence, tuber curvature, flattening and shape uniformity. The C1742-8-95/White Lady stood out as a superior cross, as did all other crosses with White Lady, regarding tuber appearance and yield traits.

Efficiency of selection in early generations of potato families with a view toward heat tolerance

The aim of this study was to evaluate the efficiency of selection of potato families in early generations for heat tolerance. Thirty families were evaluated in the seedling generation (SG), first clonal generation (FCG) in the field and greenhouse under high temperature conditions, and second clonal generation (SCG) under mild temperatures. The mean of the families was obtained in each generation. The 16 most productive families in FCG were selected, and the clones of these families were evaluated in experiments in the winter and rainy crop seasons. The results showed that family selection for tuber shape may be applied as of the SG. It was also observed that family selection in the FCG and SCG for yield and tuber specific gravity contributed to identification of clones tolerant to heat and responsive to environmental improvement.

Stable isotope ratio (¹³C/¹²C) mass spectrometry to evaluate carbon sources and sinks: changes and trends during the decomposition of vegetal debris from eucalyptus clone plantations (NW Spain)

Vegetal debris is known to participate in key soil processes such as the formation of soil organic matter (OM), also being a potential source of greenhouse gases to the atmosphere. However, its contribution to the isotopic composition of both the soil OM and the atmospheric carbon dioxide is not clear yet. Hence, the main objective of the present research is to understand the isotopic 13C changes and trends that take place during the successive biodegradative stages of decomposing soil organic inputs. By incubating bulk plant tissues for several months under laboratory controlled conditions, the kinetics of the CO2 releases and shifts in the 13C natural abundance of the solid residues were investigated using litter samples coming from forest plantations with a different clone (Anselmo: 1st clonal generation attained by morphological selection and Odiel: 2nd clonal generation genetically obtained) of Eucalyptus globulus Labill. developed over granitic or schistic bedrocks and located in northwestern Spain. Significant isotopic variations with time were observed, probably due to the isotopically heterogeneous composition of these complex substrates in conjunction with the initial selective consumption of more easily degradable 13C-differentiated compounds during the first stages of the biodegradation, while less available or recalcitrant litter components were decomposed at later stages of biodegradation, generating products that have their own specific isotopic signatures. These results, which significantly differ depending on the type of clone, suggest that caution must be exercised when interpreting carbon isotope studies (at natural abundance levels) since perturbations associated with the quality or chemical composition of the organic debris from different terrestrial ecosystems can have an important effect on the carbon stable isotope dynamics.

Selection intensities of families and clones in potato breeding

Families selection has not been recommended as a selection method for vegetative propagated species. To verify its utility for potato improvement a series of experiments were carried out under warm temperatures (rainy season). Thirty clonal families originated from heat tolerant parents were evaluated for tuber yield and specific gravity. After obtaining the seedling generation (SG) and the first clonal generation (FCG) individual clones from a further two generations were assessed. Simulations were conducted with different intensities of family selection in SG and FCG and intensities of clonal selection in subsequent generations. The results show that family selection intensities between 50% and 60% allowed the greatest gains. Estimates of h² at the families level were always higher than at the clones level and corroborate for the more effective selection of families in early generations. The selection of families for tuber specific gravity can be made in the early generations (SG and FCG) regardless of the temperature conditions where the individual clones will be selected. The sequential selection of families with intensities of 60% (SG) and 60% (FCG) would promote greater efficiency for the selection of clones in second or third clonal generation. In the case of applying stronger selection intensities for a higher efficiency with sequential selection (SG and FCG) weaker intensities should be applied in the SG and stronger intensity in FCG . Family selection for tuber yield could be practiced in the FCG as long as the selection of clones was not held in contrasting temperature conditions.

Repeatability of family means in early generations of potato under heat stress

The selection of potato clones for yield is usually effective from the second clonal generation. It would however be ideal to select from the seedling generation (SG) or the first clonal generation (FCG). The aim of this study was to evaluate the repeatability of selection at the family level performed in the early generations with the subsequent generations, for tuber yield and specific gravity in the warm season. Thirty families were evaluated in the SG, FCG and the second, third and fourth clonal generation, without any selection. In this way, adjusted means of the 30 families were obtained for each generation. Selections were simulated independently in the different family generations, and then the repeatability of these families among generations verified, using the same selection intensity. The results showed that the family selection in the SG and FCG for specific gravity and in the FCG for tuber yield is efficient, at mild as well as warm temperatures.

Early selection of full-sib potato families

Clonal selection is the preferred breeding method used in potatoes (Solanum tuberosum L.). However this selection procedure is only efficient for more advanced generations and shows no good results when applied in the seedling up to the second clonal generation. This study assessed the feasibility of selection in early generations of full-sib potato families and compares the selection method among and within families with the combined selection under different selection intensities. Six experiments were conducted from the first (C1) until the third clonal generation (C3). In C1 a randomized complete block design with four replications of 25 plants was used. In the remaining generations RCB was employed with three replications of 10 plants. Genetic variances were lower between families than within families, for all traits, but the heritabilities between families were almost always larger. The expected gains from selection between and within families were superior to gains from the combined selection in any intensity of selection. The selection of families should have weaker intensity than selection among clones within families. The selection of families was more efficient when based on the average of environments.