The relationship between fruit growth and apical senescence in the G2 line of peas

In the G2 line of peas (Pisum sativum L.), senescence of the shoot apex (which precedes leaf senescence) only occurs in long days (LD) though flowering is independent of photoperiod. It has been suggested that the photoperiodic control of senescence in G2 is mediated through different rates of seed growth. In LD seed growth is more rapid than in short days (SD) and this places a greater nutrient drain on the plant. In addition, more flowers develop into fruits in LD than in SD: 32% of flower buds abort in SD while almost none abort in LD. Senescence is associated with early seed growth and does not occur in deflowered or deseeded plants. Seed development is completed in 30d in LD while it takes 40d in SD, though the seed weights are similar. The maximum rate of fresh-weight gain of all the growing seeds of eight fruits on a plant in SD (1,440 mg/d) does not reach the maximum rate of weight gain of a similar fruit complement in LD (1,720 mg/d). The appearance of senescence symptoms in the shoot apices of LD-grown G2 plants occurs, however, prior to the time of the greatest rate of seed-weight gain. In LD, four fruits with a combined maximum growth rate of 1,250 mg/d are sufficient to cause the appearance of senescence symptoms. This is a lower combined seed growth rate than in SD where senescence does not occur. The seeds in up to 12 fruits can be growing at any time in SD with a combined maximum seed-growth rate (1,660 mg/d), only slightly less than the maximum in LD, with no sign of senescence. It is concluded that the different rates of seed growth occasioned by different photoperiods bear no relation to senescence. However, photoperiod does alter the spatial relationship of the shoot apex and the filling fruits. In LD apical growth becomes slower as fruiting proceeds so that the distance between the filling fruits and the apex is decreased to only two nodes while in SD, because of the delayed fruit development compared to LD, the spatial separation between the fruits and the shoot apex is nine nodes. Even if the growth rate of the plant had remained constant in LD it is calculated that an equivalent fruit complement would still be located three nodes further from the apex in SD than in LD. This increased spatial separation of fruits and apex in SD compared to LD probably alters the source/sink distribution of photosynthate and leaf derived hormones so that larger amounts are available to the apex in SD than LD. Also any senescence factor exported from fruits is less likely to reach the apex in SD. In continuously deflorated plants of G2 the two uppermost expanded stipules enclose the apex in SD while in LD they open out. The effect is reversible. Thus photoperiod probably affects the apex and its growth, directly, i.e. independent of fruit development, and this is accentuated by the differing spatial relationships of the apex and fruits resulting from different fruit growth rates under the different photoperiodic conditions.Abbreviations LD long day(s) - SD short day(s)     

Effect of Fruits on Dormancy and Abscisic Acid Concentration in the Axillary Buds of Phaseolus vulgaris L

The mechanism regulating the growth of adult plants in two determinate bean (Phaseolus vulgaris L.) cultivars was investigated. “Redkloud” plants flowered, formed fruits, and ceased shoot growth earlier than “Redkote” plants. Redkloud attained a smaller plant size, compared to Redkote, by imposing dormancy on axillary buds at an earlier age. In both cultivars, cessation of bud growth coincided with maximum combined fruit length per plant. Removal of fruits caused resumption of axillary bud growth within 4 to 5 days. The amount of new growth induced by fruit removal depended on the cultivar and plant age. In fully developed Redkloud plants, where shoot growth had already ceased, total leaf and shoot number per plant nearly doubled within 2 weeks following fruit removal. A much smaller response was observed in the still growing Redkote plants. Fruits, therefore, are assumed to play a major role in the regulation of shoot growth and total plant size through the control of axillary bud dormancy. It seems that smaller plant size, earlier maturity, and earlier senescence of Redkloud, compared to Redkote, were the result of earlier flowering, and accomplished in part through the growth-inhibiting action of fruits.     

BENZYLADENINE EFFFCTS ON BEAN LEAF GROWTH AND SENESCENCE

Experiments concerning the effects of benzyladenine applications on the growth and senescence of leaves have been carried out with cuttings of bean seedlings. By measuring the interactions between leaves, it was found that this kinin not only can stimulate the growth of a treated whole leaf, but it can bring about the inhibition of growth in other untreated leaves on the same plant. Consistent, with the apparent mobilizing actions of this chemical, it was found that applications to one or more leaves would induce the senescence of untreated leaves in a manner similar to the senescence-inducing effects of stem apices and flowers and fruits. The experiments suggest that the mobilization effects due to natural kinins in such centers in the intact plant may provide endogenous stimuli of leaf senescence.     

The death hormone hypothesis

The 'death hormones' or monocarpic senescence factors are hypothetical substances transported from the fruits to the vegetative parts of the mother plant, where they may stop growth, activate senescence, remobilize nutrients. and finally lead to the death of the plant. Death hormones could well be derivatives of jasmonic acid, 4-chloroindoleacetic acid, or other chlorine compounds.     

Photoperiodic and genetic control of carbon partitioning in peas and its relationship to apical senescence

Apical senescence but not flower initiation is delayed by short days (SD) compared to long days (LD) in pea plants (Pisum sativum L.) of genotype E Sn Hr. We recently reported that delay of senescence correlated with slower reproductive development, suggesting that fruits are weaker sinks for assimilates under delayed senescence conditions. Thus, we have examined assimilate partitioning in peas to determine if genotype and photoperiod regulate relative sink strength. Assimilate diversion by developing fruit has been implicated in senescence induction. A greater percentage of leaf-exported ¹⁴C was transported to fruits and a smaller percentage to the apical bud of G2 peas (genotype E Sn Hr) in LD than in SD. Relatively more of the ¹⁴C delivered to the apical bud of G2 peas was transported to flower buds than to young leaves in LD as compared to SD. There was no striking photoperiodic difference in carbon partitioning in genetic lines without the Sn Hr allele combination. The Sn Hr allele combination and photoperiod may regulate the relative strength of reproductive and vegetative sinks. Photoperiodic differences in sink strength early in reproduction suggest that these genes regulate sink strength by affecting the physiology of the whole plant. High vegetative sink strength in SD may maintain assimilate supply to the apical bud, delaying senescence.     

Overexpression of Arabidopsis hexokinase in tomato plants inhibits growth, reduces photosynthesis, and induces rapid senescence.

Sugars are key regulatory molecules that affect diverse processes in higher plants. Hexokinase is the first enzyme in hexose metabolism and may be a sugar sensor that mediates sugar regulation. We present evidence that hexokinase is involved in sensing endogenous levels of sugars in photosynthetic tissues and that it participates in the regulation of senescence, photosynthesis, and growth in seedlings as well as in mature plants. Transgenic tomato plants overexpressing the Arabidopsis hexokinase-encoding gene AtHXK1 were produced. Independent transgenic plants carrying single copies of AtHXK1 were characterized by growth inhibition, the degree of which was found to correlate directly to the expression and activity of AtHXK1. Reciprocal grafting experiments suggested that the inhibitory effect occurred when AtHXK1 was expressed in photosynthetic tissues. Accordingly, plants with increased AtHXK1 activity had reduced chlorophyll content in their leaves, reduced photosynthesis rates, and reduced photochemical quantum efficiency of photosystem II reaction centers compared with plants without increased AtHXK1 activity. In addition, the transgenic plants underwent rapid senescence, suggesting that hexokinase is also involved in senescence regulation. Fruit weight, starch content in young fruits, and total soluble solids in mature fruits were also reduced in the transgenic plants. The results indicate that endogenous hexokinase activity is not rate limiting for growth; rather, they support the role of hexokinase as a regulatory enzyme in photosynthetic tissues, in which it regulates photosynthesis, growth, and senescence.     

Programmed Cell Death in Relation to Petal Senescence in Ornamental Plants

Cell death is a common event in all types of plant organisms. Understanding the phenomenon of programmed cell death (PCD) is an important area of research for plant scientists because of its role in senescence and the post-harvest quality of ornamentals, fruits, and vegetables. In the present paper, PCD in relation to petal senescence in ornamental plants is reviewed. Morphological, anatomical, physiological,and biochemical changes that are related to PCD in petals, such as water content, sink-source relationships,hormones, genes, and signal transduction pathways, are discussed. Several approaches to improving the quality of post-harvest ornamentals are reviewed and some prospects for future research are given.     

Phytohormones and microRNAs as sensors and regulators of leaf senescence: Assigning macro roles to small molecules

Ageing or senescence is an intricate and highly synchronized developmental phase in the life of plant parts including leaf. Senescence not only means death of a plant part, but during this process, different macromolecules undergo degradation and the resulting components are transported to other parts of the plant. During the period from when a leaf is young and green to the stage when it senesces, a multitude of factors such as hormones, environmental factors and senescence associated genes (SAGs) are involved. Plant hormones including salicylic acid, abscisic acid, jasmonic acid and ethylene advance leaf senescence, whereas others like cytokinins, gibberellins, and auxins delay this process. The environmental factors which generally affect plant development and growth, can hasten senescence, the examples being nutrient dearth, water stress, pathogen attack, radiations, high temperature and light intensity, waterlogging, and air, water or soil contamination. Other important influences include carbohydrate accumulation and high carbon/nitrogen level. To date, although several genes involved in this complex process have been identified, still not much information exists in the literature on the signalling mechanism of leaf senescence. Now, the Arabidopsis mutants have paved our way and opened new vistas to elucidate the signalling mechanism of leaf senescence for which various mutants are being utilized. Recent studies demonstrating the role of microRNAs in leaf senescence have reinforced our knowledge of this intricate process. This review provides a comprehensive and critical analysis of the information gained particularly on the roles of several plant growth regulators and microRNAs in regulation of leaf senescence.     

植物一氧化氮生物学的研究进展

一氧化氮 (NO) 是植物中的一种关键的信号分子。在植物中, NO的潜在来源包括一氧化氮合成酶、硝酸还原酶、黄嘌呤氧化还原酶和非酶促途径。NO能促进植物生长, 延缓叶片、花和果实衰老, 促进休眠和需光种子的萌发, 能与植物激素相互作用调节气孔运动, 诱导程序性细胞死亡和防御相关基因的表达, 并在逆境中作为一种抗氧化剂起作用。 NO的细胞内信号反应包括环鸟苷酸、环腺苷二磷酸核糖的产生和细胞质Ca2+浓度的增加, 其信号转导途径及其生物化学和细胞学本质还不十分清楚。     

植物一氧化氮生物学的研究进展

一氧化氮(NO)是植物中的一种关键的信号分子.在植物中,NO的潜在来源包括一氧化氮合成酶、硝酸还原酶、黄嘌呤氧化还原酶和非酶促途径.NO能促进植物生长,延缓叶片、花和果实衰老,促进休眠和需光种子的萌发,能与植物激素相互作用调节气孔运动,诱导程序性细胞死亡和防御相关基因的表达,并在逆境中作为一种抗氧化剂起作用.NO的细胞内信号反应包括环鸟苷酸、环腺苷二磷酸核糖的产生和细胞质Ca2 浓度的增加,其信号转导途径及其生物化学和细胞学本质还不十分清楚.     

Programmed Cell Death in Relation to Petal Senescence in Ornamental Plants

Cell death is a common event in all types of plant organisms. Understanding the phenomenon of programmed cell death (PCD) is an important area of research for plant scientists because of its role in senescence and the post-harvest quality of ornamentals, fruits, and vegetables. In the present paper, PCD in relation to petal senescence in ornamental plants is reviewed. Morphological, anatomical, physiological,and biochemical changes that are related to PCD in petals, such as water content, sink-source relationships,hormones, genes, and signal transduction pathways, are discussed, Several approaches to improving the quality of post-harvest ornamentals are reviewed and some prospects for future research are given.     

Resource partitioning to male and female flowers of Spinacia oleracea L. in relation to whole-plant monocarpic senescence

Male plants of spinach (Spinacea oleracea L.) senesce following flowering. It has been suggested that nutrient drain by male flowers is insufficient to trigger senescence. The partitioning of radiolabelled photosynthate between vegetative and reproductive tissue was compared in male (staminate) versus female (pistillate) plants. After the start of flowering staminate plants senesce 3 weeks earlier than pistillate plants. Soon after the start of flowering, staminate plants allocated several times as much photosynthate to flowering structures as did pistillate plants. The buds of staminate flowers with developing pollen had the greatest draw of photosynthate. When the staminate plants begin to show senescence 68% of fixed C was allocated to the staminate reproductive structures. In the pistillate plants, export to the developing fruits and young flowers remained near 10% until mid-reproductive development, when it increased to 40%, declining to 27% as the plants started to senesce. These differences were also present on a sink-mass corrected basis. Flowers on staminate spinach plants develop faster than pistillate flowers and have a greater draw of photosynthate than do pistillate flowers and fruits, although for a shorter period. Pistillate plants also produce more leaf area within the inflorescence to sustain the developing fruits. The (14)C in the staminate flowers declined due to respiration, especially during pollen maturation; no such loss occurred in pistillate reproductive structures. The partitioning to the reproductive structures correlates with the greater production of floral versus vegetative tissue in staminate plants and their more rapid senescence. As at senescence the leaves still had adequate carbohydrate, the resources are clearly phloem-transported compounds other than carbohydrates. The extent of the resource redistribution to reproductive structures and away from the development of new vegetative sinks, starting very early in the reproductive phase, is sufficient to account for the triggering of senescence in the rest of the plant.     

Sucrose and Malic Acid as the Compounds Exported to the Apical Bud of Pea following CO(2) Labeling of the Fruit : No Evidence for a Senescence Factor

The G2 line of peas (Pisum sativum L.) displays senescence and death of the apical bud only in long days and in the presence of fruit. As the removal of fruit prevents senescence, one possible mechanism by which fruits induce senescence is that the fruits produce some `senescence factor' under long day conditions, which is then transported to the apical bud. Allowing developing fruits to photosynthesize in the presence of ¹⁴CO₂ results in the recovery of label in the apical bud. In order to determine the chemical nature of this radiolabeled material, fruits of G2 peas, growing under long days, were exposed to ¹⁴CO₂ at the time when the first senescence symptoms start to appear. The radiolabeled material from apical buds was then extracted, purified, and identified. Using HPLC and GC-MS the major labeled compound found in the apical bud following exposure of pea fruits to ¹⁴CO₂ was identified as sucrose, while malic acid was identified as the major ethyl acetate-soluble compound. These compounds accounted for about 73 and 16%, respectively, of the radioactivity in the apical bud. No other compounds were present in significant amounts. As neither of these chemicals is likely to have any kind of senescence effect, we report no evidence for a senescence factor.     

Modification of Seed Growth in Soybean by Physical Restraint: Effect on Leaf Senescence

Crafts-Brandner, S. J. and Egli, D. B. 1987. Modification ofseed growth in soybean by physical restraint. Effect on leafsenescence.—J. exp. Bot. 38: 2043–2049. The effect of total plant sink size on leaf senescence in soybean[Glycine max (L.) Merrill] was investigated by using a simple,non-destructive method to decrease seed growth rate and totalplant fruit sink size without altering fruit or seed number.The treatment consisted of placing plastic pod restriction devices(PPRD), which were made from plastic drinking straws (6·35mm diameter), over the fruits so that all of the seeds werecontained within the PPRD's. The treatment did not alter thetime of initiation of leaf senescence for two cultivars (McCalland Maple Amber), but decreased the rate of leaf senescencebased on declines in chlorophyll, ribulose-l,5-hi'sphosphatecarboxylase/oxygenase level and carbon dioxide exchange rate.The treatment also delayed seed maturation. At the time of seedmaturation, the plants still retained green leaves. In a separate experiment, one seed in each fruit (40% of theseeds on the plant) was not restrained by the PPRD's. This treatmentled to an intermediate rate of leaf senescence compared to controland complete seed restriction treatments. The results indicatedthat, for the cultivars examined (1) leaf senescence was initiatedat the same time regardless of sink size (2) the rate of leafsenescence could be modified by altering sink size and (3) seedmaturation could occur without complete leaf yellowing and leafabscission. The effect of the PPRD treatments on leaf senescencewere similar to results obtained when fruits were physicallyremoved, which indicated that physical removal of fruits doesnot lead to artefacts due to wounding of the plants. Key words: Glycine max L, senescence, source-sink     

Regulation by kinetin and abcisic acid of correlative senescence in relation to grain maturation,source-sink relationship and yield of rice (Oryza sativa L.)

When kinetin was applied to the source organ (flag leaf) of rice (Oryza sativa L. cv. Ratna), foliar senescence was delayed and grain yield per plant (as evidenced by grain weight, grain/straw weight ratio and 1,000 grain growth) was increased through the increase of sink activity (increase in dry weight of the grains/plant), duration of sink capacity as well as photosynthetic ability of the glumes (as determined by the chlorophyll content of the glumes of the developing grains). However, application of kinetin to the sink organs (fruits), promoted senescence of the source but increased the yield by increasing the sink capacity and 1,000 grain growth mostly at the earlier stage of reproductive development. Lower sterility percentage was associated with higher grain yield of the plant by kinetin treatments. ABA applied either to the source or the sink promoted leaf senescence and reduced the grain yield by reducing the sink activity, harvest index, sink capacity duration and increasing the sterility percentage. Thousand grain dry weight at harvest did not vary significantly amongst the treatments. It was concluded that nutrient drainage was associated with the correlative influence of fruit on the monocarpic senescence of rice plant and that a competetion for differential allocation of cytokinin and ABA in the source and sink organs initiates this senescence syndrome.