Programmed senescence of plant organs

The senescence of plant organs associated with reproductive development has been studied extensively during the past century, and it has long been recognized that this type of death is internally programmed. The regulation of organ senescence as well as its biochemical and genetic determinants has been an historically rich area of research. Certain plant hormones have been implicated as regulators or modulators of organ senescence and many of the biochemical pathways associated with the senescence syndrome have been elucidated. The genetic basis of organ senescence has also been well established by the identification of mutations that impair the senescence program and recently, transgenic plants have been used to critically determine the role of specific enzymes and hormonal signals in mediating programmed senescence of plant organs. Here, we review the current understanding of the processes that regulate leaf, flower and fruit senescence, emphasizing the role that programmed organ senescence plays in the adaptive fitness of plants.     

Effect of spikelet removal on the whole plant senescence of rice

The rice (Oryza sativa L.) cultivar IET 1444 showed a nonsequential mode of senescence as evident from the decline in chlorophyll and protein of the flag and second leaves at the senescent stage. Removal of 50,75 and 100 % spikelets from the panicle of rice plant or emasculation of the panicle by hot water treatment induced the development of secondary branch from the axil of second leaf but 25 % removal had no effect. Similarly, removal of 75 and 100 per cent spikelets from the panicle of secondary branch induced tertiary branch development, while 25 and 50 per cent removal had no such effect. Similar treatments on tertiary branch had no effect on further branch production. The pattern of leaf senescence of the untreated (control) main tiller, secondary and tertiary branches was identical, i. e. nonsequential, which could be changed into the sequential type only by the development of additional sinks (i. e. side branch). The leaf area and the seed number of secondary and tertiary branches were gradually reduced and reached a critical value in the tertiary branch. The removal of spikelets or emasculation delayed leaf senescence of the main tiller and the secondary and tertiary branches. Also the longevity of the whole plant could be increased by 40 d i. e., up to the senescence of the tertiary branch. Both leaves and reproductive parts control side branch production, which, in turn, controls the longevity of the whole rice plant.     

活性氧、自由基与植物的衰老

介绍近 1 0年来有关活性氧、自由基的产生 ,对植物的伤害及植物对活性氧、自由基清除的研究进展。     

不同植物生长调节剂对小麦衰老及产量构成的调节效应

研究了叶面喷施GA3、NBT、6－BA、TIBA、PP333对小麦旗叶衰老过程中若干生理指标的影响,分析了它们对成产要素的不同作用。结果表明：不同生长调节剂可以以不同的方式调节小麦旗叶的衰老,影响小麦产量构成因素水平。其中6－BA和PP333能够提高单位面积穗数,明显延缓叶片衰老,增加了穗粒数和千粒重,使成产3因素间比例协调合理,产量较高;TIBA可以明显提高叶绿素含量并显著提高千粒重。GA3和NBT对延缓旗叶衰老有一定作用,但增产效果不显著。研究指出,根据小麦生育状况有选择性地使用生长调节剂,协调成产因素关系,是一个有效的增产途径。     

A subtilisin-like serine protease specifically expressed in reproductive organs in rice

To identify genes specifically expressed in flowering pistils and that are related to reproductive phenomena, simplified differential display was performed with cDNA obtained from pistils and ovaries at several stages. One clone preferentially expressed in pistils at flowering and 1 day before flowering was identified as a rice subtilisin-like serine protease (RSP1). Sequence comparisons revealed that RSP1 has several characteristics in common with preproproteins. RT-PCR, northern blot, and in situ hybridization analysis revealed that RSP1 mRNA accumulates in pistils and in the filaments of stamens, whereas mRNA was undetectable in tissue from leaves, roots, panicles, and embryos. The mRNA levels in pistils increased slightly at flowering and decreased afterwards. Possible roles of the subtilisin-like serine protease in plant reproduction are discussed. Received: 27 February 2000 / Revision accepted: 24 October 2000     

Monocarpic senescence in wheat: Influence of sterile glumes and ear

The senescence of sterile glumes, flag leaf and the other two leaves below the ear of wheat ( Triticum aestivum L. cv. Sonalika) was studied in relation to grain development and surgical manipulation. The senescence of sterile glumes was faster than that of the leaves in terms of chlorophyll and protein degradation. The flag leaf senesced later than the other two leaves below it. Removal of sterile glumes markedly reduced the harvest index (crop: straw ratio) and average dry weight per grain as compared to removal of the flag leaf. Maximum grain weight was achieved after the glumes had senesced completely. Removal of the ear delayed senescence of all the three leaves. It is concluded that sterile glumes are important suppliers of assimilate for grain filling and that nutrient drainage is the primary cause of the monocarpic senescence in wheat.     

Higher salinity tolerance cultivars of winter wheat relieved senescence at reproductive stage

In order to probe into the potential of relieving senescence in the new cultivars, we have compared gas exchanges and related physiological parameters of two cultivars of winter wheat (DK961, salt-tolerant; JN17, salt-sensitive) under a series of NaCl concentrations. Through out the whole reproductive period, it was noted that net photosynthetic rate, stomatal conductance, pigment contents, ions contents, leaf area index, leaf area duration, leaf relative water content and dry matter accumulation of spikes decreased in both cultivars with saline concentrations increasing. However, the salt-tolerant cultivars showed none significant reductions in those parameters compared with control under 0.3% and 0.5% salt concentrations, with only considerable decrease happening when soil salt concentration exceeded to 0.7%. Sharply contrast to salt-tolerant cultivars, the salt-sensitive cultivars appeared linear reductions in physiological parameters under a series of salt concentrations (0.3%, 0.5% and 0.7%), with the photosynthetic duration being evidently shorter. Significantly positive correlations among K⁺/Na⁺, reproductive growth period and total growth period were noted in salt-sensitive cultivars, however, none significant relations appeared among those parameters in salt-tolerant cultivars, indicating a strong tolerant behavior happened. Our result suggested that higher salinity tolerance cultivars of winter wheat could relieve senescence at the reproductive stage.     

Effect of respiratory homeostasis on plant growth in cultivars of wheat and rice

Some plants have the ability to maintain similar respiratory rates (measured at the growth temperature), even when grown at different temperatures, a phenomenon referred to as respiratory homeostasis. The underlying mechanisms and ecological importance of this respiratory homeostasis are not understood. In order to understand this, root respiration and plant growth were investigated in two wheat cultivars (Triticum aestivum L. cv. Stiletto and cv. Patterson) with a high degree of homeostasis, and in one wheat cultivar (T. aestivum L. cv. Brookton) and one rice cultivar (Oryza sativa L. cv. Amaroo) with a low degree of homeostasis. The degree of homeostasis (H) is defined as a quantitative value, which occurs between 0 (no acclimation) and 1 (full acclimation). These plants were grown hydroponically at constant 15 or 25 °C. A good correlation was observed between the rate of root respiration and the relative growth rates (RGR) of whole plant, shoot or root. The plants with high H showed a tendency to maintain their RGR, irrespective of growth temperature, whereas the plants with low H grown at 15 °C showed lower RGR than those grown at 25 °C. Among several parameters of growth analysis, variation in net assimilation rate per shoot mass (NAR_m) appeared to be responsible for the variation in RGR and rates of root respiration in the four cultivars. The plants with high H maintained their NAR_m at low growth temperature, but the plants with low H grown at 15 °C showed lower NAR_m than those grown at 25 °C. It is concluded that respiratory homeostasis in roots would help to maintain growth rate at low temperature due to a smaller decrease in net carbon gain at low temperature. Alternatively, growth rate per se may control the demand of respiratory ATP, root respiration rates and sink demands of photosynthesis. The contribution of nitrogen uptake to total respiratory costs was also estimated, and the effects of a nitrogen leak out of the roots and the efficiency of respiration on those costs are discussed.     

Influence of diamines and polyamines on the senescence of plant suspension cultures

The diamines putrescine and cadaverine and the polyamines spermine and spermidine inhibited the senescence of nonphotosynthetic cultures of Paul's Scarlet rose. Response was observed when the media of stationary phase cultures was adjusted to either 1 mM of cadaverine or putrescine; or 0.1 μM of either spermine or spermidine along with 2% sucrose in all cases. Senescence of the cultures was followed by microscopic examination of cell aliquots removed at 10 day intervals and treated with the vital stain, fluorescein diacetate.     

Variation in economically and ecologically important traits in onion plant organs during reproductive development

The spatial distribution of organosulphur compounds throughout the onion (Allium cepa L.) plant body during reproduction is of ecological and horticultural interest. These secondary metabolites are associated with both pest resistance and many of the vegetable's culinary and medicinal properties, including the ability to inhibit platelet aggregation. Inhibition of platelet aggregation can be of benefit to human cardiovascular health. Organosulphur compound concentrations are associated with elemental sulphur, pungency, soluble solids and effect on human platelet aggregation. These parameters were evaluated in extracts collected separately from bulb scales, leaf blades, scapes and umbels biweekly throughout the reproductive phase of the life cycle of the onion. Significant variation in pungency, platelet inhibition, total sulphur content and soluble solids existed among samples of organs and within organs over time during reproductive growth. Furthermore, some extracts from leaf, scape and bulb induced rather than inhibited platelet aggregation.     

Role of other plant organs in gibberellic acid-induced delay of leaf senescence in alstroemeria cut flowers

Chlorophyll loss in leaves of cut flowers of alstroemeria (Alstroemeria pelegrina L. cv. Westland) was rapid in darkness and counteracted by irradiation and treatment of the flowers with gibberellic acid (GA₃). The mechanism of the effect of GA₃ under dark conditions was investigated. The content of various carbohydrates in the leaves under dark conditions rapidly decreased; this was not influenced by treatment with GA_3. indicating that the loss of carbohydrates in the leaves did not induce the loss of chlorophyll. Placing the cut flowers in various solutions of organic and inorganic nutrients exhibited no significant effect on the retention of chlorophyll in leaves of dark-senescing flowers. The total nitrogen content in leaves of dark-senescing cut flowers decreased with time. Leaves of GA₃-treated flowers retained more nitrogen. In contrast, the buds of GA₃-treated flowers retained less nitrogen during senescence in the dark than control buds. To investigate whether GA₃ affects export of assimilates from the leaf to various parts of control and GA₃-treated flowers, we labelled one leaf with radioactive carbon dioxide. ¹⁴C-assimilates accumulated preferentially in the flowers, in which the relative specific activity of the youngest floral buds was highest. No significant differences were observed in the distribution of ¹⁴C-labelled compounds between the buds of control and GA₃-treated flowers. To establish the importance of source-sink relations for the loss of leaf chlorophyll we removed the flower buds (i. e. the strongest sink) from the cut flowers. This removal only slightly delayed chlorophyll loss as compared to the large delay caused by GA₃-treatment. In addition, detached leaf tips exhibited chlorophyll loss in the dark, which was delayed by GA₃-treatment in a fashion comparable with that in flowers. Together these data demonstrate that interactions of the leaves with other plant organs are not essential for chlorophyll loss during senescence in the dark. Additionally, we have found no evidence that GA₃ delays the loss of chlorophyll by affecting the transport of nutrients within the cut flowers.     

Ephemerality and reproductive senescence in avian filarioids

Adult Eulimdana spp (Nemotoda, Filarioidea) of shorebirds (Charodrii formes) produce long-lived, skin-inhabiting microfilariae and then die and are resorbed. The host is thereafter refractory to infection. Pelecitus fulicaeatrae of coots (Fulica americana) also produces long lived, skin-inhabitingmicro filariae, but adult worms become reproductively senescent. Ephemerality of adults in Eulimdana spp and reproductive senescence in P. fulicaeatrae ensure that the skin will not become saturated with microfilariae, large numbers of which would probably be harmful to mallophogan vectors that remain on the host and are continually exposed to micro filariae while feeding. Here Roy C. Anderson and Cheryl M. Bartlett discuss the significance of ephemerality and senescence in the life histories o f avian f filarioids transmitted by permanent ectoparasites.     

Glycollate metabolism of wheat and rice leaves during senescence and under the influence of growth regulators

The glycollate metabolism of wheat (Triticum vulgare Vill. cv. Sonalika) and rice (Oryza sativa L. ev. Jaya) leaves was studied during senescence by estimating the endogenous levels of glycollate and hydrogen peroxide (H₂O₂) and the activities of glycollate oxidase and catalase. In comparison with light incubation the incubation of excised leaves in the dark caused a decline in the glycollate content and in the activities of glycollate oxidase and catalase, and an increase in the H₂O₂ content, more marked in the leaves of rice than in the leaves of wheat. Glycollate oxidase activity gradually decreased with incubation time, and glycollate metabolism decreased during senescence. The glycollate oxidase in particular and glycollate metabolism of rice were more sensitive to incubation time than those of wheat. Kinetin increased the glycollate oxidase activity and glycollate metabolism during senescence, while ethrel (2-chloroethylpho-sphonic acid) and ABA (abscisic acid) reduced these activities in both plant species.     

施肥深度对旱地小麦花后根系衰老的影响

研究了施肥深度对旱地小麦花后根系衰老的影响,结果层次（20－40cm)施肥,其根中保护酶系SOD,CAT活性可保持较高水平,根系活力和根可溶性蛋白质含量降低慢,抑制膜脂过氧化产物MDA的产生,利于延迟根系衰老,施肥过浅（0－20cm)和施肥过深（60－80cm）,根系衰老快,产量较低,从整体看,处理产生,利于延迟根系衰耆 ,施肥过浅（0－20cm施肥）及处理4（60－80cm施肥）差异显著,与处理3（40－60cm施肥）差异较少,可以认为,延迟旱地小麦根系衰老的施肥深度应在20－40cm左右,据此制定管理方案,以获得旱地小麦的高产高效。     

Age-specific mating strategies and reproductive senescence

While males gain obvious direct advantages from multiple mating, the reproductive capacity of females is more constrained. The reason why polyandry evolved in females is therefore open to many conjectures. One hypothesis postulates that females gain indirect benefits by increasing the probability of siring young from high quality males. To explore this hypothesis, we used the natural variation of the reproductive value that males and females undergo through age. The age-related variation of phenotypic performance might then induce variations in mating strategies in males and females. Using the common lizard (Lacerta vivipara) as our model system, we showed that reproductive immaturity and senescence created variability in both male and female reproductive success (including survival of offspring). Consistent with theory, males at their best-performing phenotype adopted a polygynous strategy. These males were of an intermediate age and they produced offspring of higher viability than younger and older males. In contrast, females at their best performing phenotype, also of an intermediate age, were less polyandrous than other less-performing females. Middle-aged females tended to mate with males of an intermediate age and produced litters with higher viability independently from their reproductive strategy. Males of an intermediate age enhanced their fitness by additional matings with young or old females. Young and old females increased their fitness by being more polyandrous. Polyandry therefore appears as means to seek for good males. A positive correlation between males and their partners' fitness disagree with the idea that polyandry is the result of a sexual conflict in this species.     

Influence of plant reproductive systems on the evolution of hummingbird pollination

Many hummingbird‐pollinated plant species evolved from bee‐pollinated ancestors independently in many different habitats in North and South America. The mechanisms leading to these transitions are not completely understood. We conducted pollination and germination experiments and analyzed additional reproductive traits in three sister species pairs of which one species is bee‐ and the other hummingbird‐pollinated. All hummingbird‐pollinated species showed higher seed set and germination rates in cross‐pollinated than in self‐pollinated flowers. In the self‐compatible, bee‐pollinated sister species this difference did not exist. As expected, seed set and germination rate were higher after cross‐pollination in the largely self‐incompatible genus Penstemon independently of the pollination syndrome. However, the bird‐pollinated species produce only half of the amount of ovules and pollen grains per flower compared to the bee‐pollinated sister species. This indicates that hummingbird pollination is much more efficient in self‐incompatible populations because hummingbirds waste less pollen and provide higher outcrossing rates. Therefore, hummingbird pollination is less resource costly. Overall, we suggest that hummingbirds may increase the reproductive success compared to bees, influencing the evolution of hummingbird pollination in ecosystems with diverse bee assemblages.     

Mechanism of monocarpic senescence in rice

During grain formation stage (90 to 110 days), the youngest flag leaf of rice (Oryza sativa L. cv. Jaya) remained metabolically most active (as indicated by cellular constituents and enzyme activities) and the third leaf the least active. At the grain development stage (110 to 120 days) the above pattern of age-related senescence of the flag leaf completely changed and it senesced at a faster rate than the second leaf which remained metabolically active even up to grain maturation time (120 to 130 days), when both the flag and the third leaf partially senesced. Removal of any leaf temporarily arrested senescence of the remaining attached leaves, that of flag leaf did not hasten senescence of the second leaf, while that of either the second or the third accelerated senescence of the flag. Removal of the inflorescence after emergence or foliar treatment of intact plant with kinetin equally delayed senescence and produced an age-related, sequential mode of senescence or leaves. Both translocation and retention of ³²P by the flag leaf were maximum at the time of grain formation and that by the second leaf was maintained even up to grain maturation time. The induction of senescence of the flag leaf was preceded by a plentiful transport of ³²P to the grains. Kinetin treatment decreased the transport of ³²P, prolonged its duration, and almost equally involved all of the leaves in this process. The pattern of senescence of isolated leaf tips was similar to that of attached leaves. The level of endogenous abscisic acid-like substance(s) maintained a close linearity with the senescence behavior of the leaves of intact and defruited plants during aging, and the rise in abscisic acid in the flag leaf was also preceded by higher ³²P transport to the grains.