1 – The Phenomena of Senescence and Aging

Publisher Summary This chapter discusses causes of monocarpic senescence and senescence in polycarpic plants and clones. Whole plant senescence is viewed primarily in terms of leaf senescence and it is measured mainly through chloroplastic parameters such as chlorophyll loss and decreased photosynthesis. In monocarpic plants where senescence and death closely follow reproductive development, the senescence (monocarpic senescence) is often controlled by the developing reproductive structures. In these cases, removal of the reproductive structures or prevention of their development usually prolongs the life of the plant. Monocarpic plants cease their vegetative growth fairly abruptly early in their reproductive phase. Conversely, the perennial polycarpic pattern requires continued vegetative growth. This prominent shift (diversion) in growth-related allocation of resources in monocarpy seems to be part of a reproductive strategy that optimizes reproductive output for the plants. This diversion/withdrawal is often quite prominent that leads to monocarpic senescence.

Whole plant senescence

Programmed cell death (apoptosis) is well-established in many multicellular organisms. Apoptosis purifies a tissue from cells that became useless or even harmful for the organism. Similar phenomena are already described also at subcellular level (suicide of mitochondria, i.e., mitoptosis) as well as at supracellular level (degradation of some organs temporarily appearing in the course of ontogenesis and then disappearing by means of apoptosis of the organ-composing cells). Following the same logic, one may put a question about programmed death of an organism as a mechanism of purification of a kin, community of organisms, or population from individuals who became unwanted for this kin, etc. A putative mechanism of such kind is proposed to be coined "phenoptosis" by analogy with apoptosis and mitoptosis. In a unicellular organism (the bacterium Escherichia coli), three different biochemical mechanisms of programmed death are identified. All of them are actuated by the appearance of phages inside the bacterial cell. This may be regarded as a precedent of phenoptosis which prevents expansion of the phage infection among E. coli cells. Purification of a population from infected individuals looks like an evolutionary invention useful for a species. Such an invention has high chances to be also employed by multicellular organisms. Most probably, septic shock in animals and humans serves as an analog of the phage-induced bacterial phenoptosis. It is hypothesized that the stress-induced ischemic diseases of brain and heart as well as carcinogenesis if they are induced by repeated stresses also represent phenoptoses that, in contrast to sepsis, are age-dependent. There are interrelations of programmed death phenomena at various levels of complexity of the living systems. Thus, extensive mitoptosis in a cell leads to apoptotic death of this cell and extensive apoptosis in an organ of vital importance results in phenoptotic death of an individual. In line with this logic, some cases are already described when inhibition of apoptosis strongly improves the postischemic state of the organism.

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Phenoptosis: Programmed Death of an Organism

9 – Cytokinins and Senescence

Cytokinins (CKs) coming from the roots via the xylem are known to delay leaf senescence, and their decline may be important in the senescence of soybean (Glycine max) plants during pod development (monocarpic senescence). Therefore, using radioimmunoassay of highly purified CKs, we quantified the zeatin (Z), zeatin riboside (ZR), the dihydro derivatives (DZ, DZR), the O-glucosides, and DZ nucleotide in xylem sap collected from root stocks under pressure at various stages of pod development. Z, ZR, DZ, and DZR dropped sharply during early pod development to levels below those expected to retard senescence. Pod removal at full extension, which delayed leaf senescence, caused an increase in xylem sap CKs (particularly ZR and DZR), while depodding at late podfill, which did not delay senescence, likewise did not increase the CK levels greatly. The levels of the O-glucosides and the DZ nucleotide were relatively low, and they showed less change with senescence or depodding. The differences in the responses of individual CKs to senescence and depodding suggest differences in their metabolism. Judging from their activity, concentrations and response to depodding, DZR and ZR may be the most important senescence retardants in soybean xylem sap. These data also suggest that the pods can depress CK production by the roots at an early stage and this decrease in CK production is required for monocarpic senescence in soybean.

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Correlation of xylem sap cytokinin levels with monocarpic senescence in soybean.

Excised soybean (Glycine max [L] Merrill) cv Anoka leaf discs tend to remain green even after the corresponding intact leaves have turned yello on fruiting plants We have found that explants which include a leaf along with a stem segment (below the node) and one or more pods (maintained on distilled H(2)O) show similar but accelerated leaf yellowing and abscission compared with intact plants In podded explants excised at pre-podfill, the leaves begin to yellow after 16 days, whereas those excised at late podfill begin to yellow after only 6 days Although stomatal resistances remain low during the first light period after excision, they subsequently increase to levels above those in leaves of intact plants Explants taken at mid to late podfill with one or more pods per node behave like intact plants in that pod load does not affect the time lag to leaf yellowing Explant leaf yellowing and abscission are delayed by removal of the pods or seeds or by incubation in complete mineral nutrient solution or in 46 micromolar zeatin Like chorophyll breakdown, protein loss is accelerated in the explants, but minerals or especially zeatin can retard the loss Pods on explants show rates and patterns of color change (green to yellow to brown) similar to those of pods on intact plants These changes start earlier in explants on water than in intact plants, but they can be delayed by adding zeatin Seed dry weight increased in explants, almost as much as in intact plants Explants appear to be good analogs of the corresponding parts of the intact plant, and they should prove useful for analyzing pod development and mechanisms of foliar senescence Moreover, our data suggest that the flux of minerals and cytokinin from the roots could influence foliar senescence in soybeans, but increased stomatal resistance does not seem to cause foliar senescence

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Anthony T. Tucker

Papers

Characterization of leaf senescence and pod development in soybean explants.