Leaves provide energy for plants, and consequently for animals, through photosynthesis. Despite their important functions, plant leaf developmental processes and their underlying mechanisms have not been well characterized. Here, we provide a holistic description of leaf developmental processes that is centered on cytokinins and their signaling functions. Cytokinins maintain the growth potential (pluripotency) of shoot apical meristems, which provide stem cells for the generation of leaf primordia during the initial stage of leaf formation; cytokinins and auxins, as well as their interaction, determine the phyllotaxis pattern. The activities of cytokinins in various regions of the leaf, especially at the margins, collectively determine the final leaf morphology (e.g., simple or compound). The area of a leaf is generally determined by the number and size of the cells in the leaf. Cytokinins promote cell division and increase cell expansion during the proliferation and expansion stages of leaf cell development, respectively. During leaf senescence, cytokinins reduce sugar accumulation, increase chlorophyll synthesis, and prolong the leaf photosynthetic period. We also briefly describe the roles of other hormones, including auxin and ethylene, during the whole leaf developmental process. In this study, we review the regulatory roles of cytokinins in various leaf developmental stages, with a focus on cytokinin metabolism and signal transduction processes, in order to shed light on the molecular mechanisms underlying leaf development.
Temperature change is of potential to trigger the formation of unreduced gametes. In this study, we showed that short periods of high temperature treatment can induce the production of 2n pollen in Populus pseudo-simonii Kitag. The meiotic stage, duration of treatment, and temperature have significant effects on the induction of 2n pollen. Heat stress resulted in meiotic abnormalities, including failure of chromosome separation, chromosome stickiness, laggards and micronuclei. Spindle disorientations in the second meiotic division, such as parallel, fused, and tripolar spindles, either increased in frequency or were induced de novo by high temperature treatment. We found that the high temperature treatment induced depolymerisation of meiotic microtubular cytoskeleton, resulting in the failure of chromosome segregation. New microtubular cytoskeletons were able to repolymerise in some heat-treated cells after transferring them to normal conditions. However, aberrant cytokinesis occurred owing to defects of new radial microtubule systems, leading to production of monads, dyads, triads, and polyads. This suggested that depolymerisation and incomplete restoration of microtubules may be important for high temperature-induction of unreduced gametes. These findings might help us understand how polyploidisation is induced by temperature-related stress and support the potential effects of global climate change on reproductive development of plants.
Tetraploid plants were produced from leaf explants of diploid Populus pseudo-simonii by treating the leaves with colchicine. Leaf explants were cultured on MS basal medium containing 1.78 μM BA and 1.08 μM NAA for 0, 6 and 12 days, and then transferred to the same MS liquid medium with colchicine at concentrations of 25, 50 and 75 μM for 1, 2 and 3 days. The highest efficiency of tetraploid induction was 14.6% by treating leaf explants that were pre-cultured for 6 days and then cultured in liquid MS with 50 μM colchicine for 3 days. Flow cytometric analysis was used to screen the tetraploids out from the regenerated plants and chromosome number counting was employed to confirm the polyploidy level. Size and frequency of leaf stomata between diploid and tetraploid plants were demonstrated to have significant differences.
Diploid (2n) eggs were induced by treating developing embryo sacs of Populus with colchicine solution, in order to produce triploid plants. The optimal pollinated time of female catkins was confirmed as timing point for each treatment. When female catkins of P. pseudo-simonii x P. nigra ‘Zheyin3#’ had become 5.62 ± 0.13 cm long 84 h after they emerged from their bract scales and all stigmas were exposed, pistils all over the entire catkin had optimal stigma receptivity. Observation of paraffin sections showed that embryo sac development of ‘Zheyin3#’, which initiated 12 h before pollination and finished 132 h after pollination, was a successive and asynchronous process. Generative cell division of pollen of the male parent P. x beijingensis took place 3-16 h after pollination. Catkins of 18-96 h after pollination of ‘Zheyin3#’ were treated with colchicine solution. In the progeny, twenty three triploids were detected by chromosome counting and the highest rate of triploids was 66.7% in one treatment. The rate of triploid yield was positively correlated with the frequency of four-nucleate embryo sacs (r = 0.6721, p = 0.0981) and was not significantly correlated with the percentages of uni-, twoand eight-nucleate embryo sac (r = -0.1667, p = 0.7210, r = -0.3069, p = 0.5031 and r = 0.0189, p = 0.9679, respectively), suggesting that the third mitotic division of embryo sac may be the effective stage to induce 2n eggs. Through this approach, completely homozygous 2n eggs can be produced. Its significance for plant breeding is discussed.
Summary Although polyploid plants have larger leaves than their diploid counterparts, the molecular mechanisms underlying this difference (or trait) remain elusive. Differentially expressed genes (DEGs) between triploid and full‐sib diploid poplar trees were identified from two transcriptomic data sets followed by a gene association study among DEGs to identify key leaf growth regulators. Yeast one‐hybrid system, electrophoretic mobility shift assay, and dual‐luciferase assay were employed to substantiate that PpnGRF5‐1 directly regulated PpnCKX1. The interactions between PpnGRF5‐1 and growth‐regulating factor (GRF)‐interacting factors (GIFs) were experimentally validated and a multilayered hierarchical regulatory network (ML‐hGRN)‐mediated by PpnGRF5‐1 was constructed with top‐down graphic Gaussian model (GGM) algorithm by combining RNA‐sequencing data from its overexpression lines and DAP‐sequencing data. PpnGRF5‐1 is a negative regulator of PpnCKX1. Overexpression of PpnGRF5‐1 in diploid transgenic lines resulted in larger leaves resembling those of triploids, and significantly increased zeatin and isopentenyladenine in the apical buds and third leaves. PpnGRF5‐1 also interacted with GIFs to increase its regulatory diversity and capacity. An ML‐hGRN‐mediated by PpnGRF5‐1 was obtained and could largely elucidate larger leaves. PpnGRF5‐1 and the ML‐hGRN‐mediated by PpnGRF5‐1 were underlying the leaf growth and development.
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