It emerged recently that there is an inter-relationship between drought and ultraviolet-B (UV-B) radiation in plant responses, in that both stresses provoke an oxidative burst. The purpose of this investigation was to compare the effects and interaction of drought and UV-B in wheat and pea. The absence of changes in relative leaf water content (RWC) after UV-B treatments indicate that changes in water content were not involved. RWC was the main factor resulting in reduced growth in response to drought. Increases in anthocyanin and phenols were detected after exposure to UV-B. The increases do not appear to be of sufficient magnitude to act as a UV-B screen. UV-B application caused greater membrane damage than drought stress, as assessed by lipid peroxidation as well as osmolyte leakage. An increase in the specific activities of antioxidant enzymes was measured after UV-B alone as well as after application to droughted plants. Proline increased primarily in drought-stressed pea or wheat. Proline may be the drought-induced factor which has a protective role in response to UV-B. The physiological and biochemical parameters measured indicate the UV-B light has stronger stress effectors than drought on the growth of seedlings of both species. The two environmental stresses acted synergistically to induce protective mechanisms in that pre-application of either stress reduced the damage caused by subsequent application of the other stress.

https://onlinelibrary.wiley.com/doi/pdf/10.1046/j.1365-3040.2001.00778.x

The effect of drought and ultraviolet radiation on growth and stress markers in pea and wheat

SUMMARY
Leaf senescence is a hiphly-controlled sequence of events comprising the final stage of development. Cells remain viable during the process and new gene expression is required. There is some similarity between senescence in plants and programmed cell death in animals. In this review, different classes of senescence-related genes are defined and progress towards isolating such genes is reported. A range of internal and external factors which appear to cause leaf senescence is considered and various models for the mechanism of senescence- initiation are described. The current understanding of senescence at the wrganelle and molecular levels is presented. Finally, same ideas are mooted as to why senescence occurs and why it should be studied further.

Gene expression during leaf senescence

(1988). Inhibition of legume nodule formation and N2 fixation by nitrate. Critical Reviews in Plant Sciences: Vol. 7, No. 1, pp. 1-23.

Inhibition of legume nodule formation and N2 fixation by nitrate

Factors that influence the longevity and senescence of photosynthetic tissues of Arabidopsis were investigated. To determine the influence of reproductive development on the timing of somatic tissue senescence, the longevity of rosette leaves of the Landsberg erecta strain and of isogenic mutant lines in which flowering is delayed (co-2) or sterile flowers are produced (ms1-1) were compared. No difference in the timing of senescence of individual leaves was observed between these lines, indicating that somatic tissue longevity is not governed by reproductive development in this species. To examine the role of differential gene expression in the process of leaf senescence, cDNA clones representing genes that are differentially expressed in senescing tissues were isolated. Sequence analysis of one such clone indicated homology to previously cloned cysteine proteinases, which is consistent with a role for the product of this gene in nitrogen salvage. RNA gel blot analysis revealed that increased expression of senescence-associated genes is preceded by declines in photosynthesis and in the expression of photosynthesis-associated genes. A model is presented in which it is postulated that leaf senescence is triggered by age-related declines in photosynthetic processes.

Developmental and age-related processes that influence the longevity and senescence of photosynthetic tissues in arabidopsis.

The partitioning of nitrate assimilation between root and shoot of higher plant species is indicated by the relative proportions of total plant nitrate reductase activity (NRA) in the two plant parts and the relative concentrations of nitrate and reduced N in the xylem sap. These have been collated here from the literature and temperate and tropical species compared.

Both the distribution of NRA and xylem sap nitrate: reduced N indicate that the following four generalizations can be made.

1Temperate, perennial species growing in low external nitrate concentrations (about 1 mol m−3) carry out most of their nitrate assimilation in the root. As external nitrate concentration increases (in the range found in agricultural soils, 1–20 mol m−3), shoot nitrate assimilation becomes increasingly important.
2Temperate, annual legume species growing in low external nitrate concentrations carry out most of their nitrate assimilation in the root. Shoot nitrate assimilation increases in importance as external nitrate concentration is increased.
3Temperate, annual non-legume species vary greatly in their partitioning of nitrate assimilation between root and shoot when growing in low external nitrate concentrations. Regardless of the proportion carried out in the root at low external nitrate concentrations, nitrate assimilation in the shoot becomes increasingly important as external nitrate concentration is increased.
4Tropical and subtropical species, annual and perennial, carry out a substantial proportion of their nitrate assimilation in the shoot when growing in low external nitrate concentrations. The partitioning of nitrate assimilation between root and shoot remains constant as external nitrate concentration increases.


It is proposed that a greater proportion of nitrate assimilation occurs in the shoot when an increase in the rate of nitrate uptake does not induce an increase in NR level in the root. Thus, a greater proportion of the nitrate taken up remains unassimilated and is passed into the xylem. A constant partitioning of nitrate assimilation between root and shoot is achieved by balancing NR levels in the root with rates of nitrate uptake.

The advantages and disadvantages of assimilating nitrate in either the root or shoot are discussed in relation to temperate and tropical habitats.

The partitioning of nitrate assimilation between root and shoot of higher plants

Fertilizer N availability impacts photosynthesis and crop performance, although cause–effect relationships are not well established, especially for field-grown plants. Our objective was to determine the relationship between N supply and photosynthetic capacity estimated by leaf area index (LAI) and single leaf photosynthesis using genetically diverse field-grown maize (Zea mays L.) hybrids. We compared a high yield potential commercial hybrid (FR1064 x LH185) and an experimental hybrid (FR1064 x IHP) with low yield potential but exceptionally high grain protein concentration. Plant biomass and physiological traits were measured at tassel emergence (VT) and at the grain milk stage (R3) to assess the effects of N supply on photosynthetic source capacity and N uptake, and grain yield and grain N were measured at maturity. Grain yield of FR1064 x LH185 was much greater than FR1064 x IHP even though plant biomass and LAI were larger for FR1064 x IHP, and single leaf photosynthesis was similar for both hybrids. Although photosynthetic capacity was not related to hybrid differences in productivity, increasing N supply led to proportional increases in grain yield, plant biomass, LAI, photosynthesis, and Rubisco and PEP carboxylase activities for both hybrids. Thus, a positive relationship between photosynthetic capacity and yield was revealed by hybrid response to N supply, and the relationship was similar for hybrids with a marked difference in yield potential. For both hybrids the N response of single leaf CER and initial Rubisco activity was negative when expressed per unit of leaf N. In contrast, PEP carboxylase activity per unit leaf N increased in response to N availability, indicating that PEP carboxylase served as a reservoir for excess N accumulation in field-grown maize leaves. The correlation between CER and initial Rubisco activity was highly significant when expressed on a leaf area or a total leaf basis. The results suggest that regardless of genotypic yield potential, maize CER, and potentially grain yield, could be improved by increasing the partitioning of N into Rubisco.

https://link.springer.com/content/pdf/10.1007%2Fs11104-008-9767-1.pdf

Physiological N response of field-grown maize hybrids (Zea mays L.) with divergent yield potential and grain protein concentration.

Visual senescence symptoms and associated changes in constituent contents of three field-grown maize ( Zea mays L.) hybrids (Pioneer brand 3382, B73 × Mo17, and Farm Service brand 854) were compared in response to ear removal. Whole plants were harvested at eight intervals during the grain-filling period, and analyzed for dry matter, total N and nitrate N, phosphorus, sugars, and starch. Upper leaves of earless P3382 and B73 × Mo17 showed reddish discoloration by 25 days after anthesis (DAA) and all leaves had lost most of their chlorophyll by 40 DAA. In striking contrast, leaves of earless FS854 plants remained green and similar in appearance to eared controls throughout the grain-filling period. For all hybrids, ear removal led to a decrease in dry weight, reduced N, total N, and phosphorus contents of the total plant, and an increase in carbohydrate content of the leaves and stalks, relative to respective controls. Although changes in carbohydrate and N contents, which previously had been associated with senescence, were observed for all earless hybrids, these changes were followed by accelerated senescence and early death only for P3382 and B73 × Mo17. By 30 DAA, earless P3382 and B73 × Mo17 plants ceased to accumulate dry weight, total N, and phosphorus, indicating a termination of major metabolic activities. In contrast, earless FS854 plants retained a portion of these metabolic activities until 58 DAA, indicating a role for roots in determining rate of senescence development. Thus, the course of senescence was more accurately reflected by measurements of metabolic activities than by measurements of metabolite contents at any given time. These results show that the ear per se does not dictate the rate or completion of the senescence process, and implicated an association between the continued accumulation of N and associated root activities with the delayed senescence pattern of the earless FS854 plants. It is evident that studies involving control of senescence among species must also consider genotypic influences within species.

https://www.jstor.org/stable/pdfplus/4268466.pdf

Differential Senescence of Maize Hybrids following Ear Removal : I. Whole Plant.

In conjunction with a study of the effects of ear removal on the senescence of whole maize ( Zea mays L.) plants, visual symptoms and associated changes in constituent contents and activities of a selected leaf (first leaf above the ear) were determined. Leaves were sampled from field-grown eared and earless Pioneer brand 3382, B73 × Mo17, and Farm Services brand 854 maize hybrids at nine times during the grainfilling period. Visual symptoms indicated the following sequence and rate of senescence: earless B73 × Mo17 > earless P3382 » eared B73 × Mo17 » eared P3382 ≤ earless FS854 > eared FS854. All earless hybrids showed increases in leaf dry weight and sugar content; however, the increases were transitory for P3382 and B73 × Mo17, but continuous throughout the grain-filling period for FS854, indicative of continued photosynthetic activity of the latter. All earless hybrids exhibited similar and transitory starch accumulation patterns. Thus, FS854 was an exception to the concept that carbohydrate accumulation accelerates leaf senescence. Ear removal resulted in accelerated losses of reduced N, phosphoenolpyruvate and ribulose bisphosphate carboxylases, phosphorus, chlorophyll, nitrate reductase activity, and moisture for P3382 and B73 × Mo17 plants. In contrast, the loss of all components (except phosphorus) was similar for the selected leaf of earless and eared FS854. Although the loss of nitrate reductase activity, reduced N, and carboxylating enzymes accurately reflected the development of senescence of the selected leaf, the rate of net loss of reduced N and carboxylating enzymes appeared to be regulated. We deduced that the rate of flux of N into the leaf was a factor in regulating the differing rates of senescence observed for the six treatments; however, we cannot rule out the possibility of concurrent influence of growth regulators or other metabolites.

https://www.jstor.org/stable/pdfplus/4268467.pdf

Differential Senescence of Maize Hybrids following Ear Removal II. Selected Leaf

The objectives of this work were to determine the effect of sink strength (presence or absence of pods) and nitrogen source (nodulating versus nonnodulating plants) on enzymic activities, chlorophyll concentration, and senescence of soybean ( Glycine max [L] Merr cv Harosoy) isolines A 2-year (1981-1982) field study was conducted For both nodulated and nonnodulated plants, ribulose bisphosphate carboxylase (RuBPCase) activity of upper-canopy leaves was decreased by pod removal in both years, while chlorophyll concentration was decreased in 1981 only Nonnodulated plants had lower RuBPCase activity in 1981 and lower chlorophyll concentration in both years compared with nodulated plants In both years, and for all treatments, RuBPCase activity and chlorophyll began to decline at about the same time, but the rate of decline was less for depodded than for podded plants Leaves in the middle and lower parts of the canopy had similar RuBPCase activity and chlorophyll concentration trends as upper-canopy leaves for all treatments Profiles of nitrate reductase activity (NRA) were similar for all treatments in both 1981 and 1982 Acetylene reduction profiles were similar for nodulated-podded and nodulated-depodded plants The peak and decline in NRA profiles preceded the peak and decline in acetylene reduction profiles The rate of decline in acetylene reduction activity was less for depodded plants, especially in 1982, but activities reached zero by the final sampling time Thus, nodule senescence was not prevented by pod removal Based on seasonal profiles of RuBPCase activity, chlorophyll, NRA, and acetylene reduction activity, the initiation of senescence appeared to occur at the same approximate time for all treatments and, thus, did not depend on the presence or absence of pods or nodules The hypothesis that nodules act as a nitrogen source and carbohydrate sink to delay senescence in the absence of pods was not correct

Effects of Pod Removal on Metabolism and Senescence of Nodulating and Nonnodulating Soybean Isolines: II. Enzymes and Chlorophyll.

Studies were conducted with 9 to 12 day-old soybean (Glycine max [L.] Merr. cv. Williams) seedlings to determine the contribution of roots to whole plant NO(3) (-) reduction. Using an in vivo -NO(3) (-) nitrate reductase (NR) assay (no exogenous NO(3) (-) added to incubation medium) developed for roots, the roots accounted for approximately 30% of whole plant nitrate reductase activity (NRA) of plants grown on 15 mm NO(3) (-).Nitrogen analyses of xylem exudate showed that 53 to 66% of the total-N was as reduced-N, depending on the time of day of exudate collection. These observations supported enzyme data that suggested roots were contributing significantly to whole plant NO(3) (-) reduction. In short-term feeding studies using (15)N-NO(3) (-) significant and increasing atom percent (15)N excess was found in the reduced-N fraction of xylem exudate at 1.5 and 3 hours after feeding, respectively, which verified that roots were capable of reducing NO(3) (-).Estimated reduced-N accumulation by plants based on in vivo -NO(3) (-) NR assays of all plant parts substantially over-estimated actual reduced-N accumulation by the plants. Thus, the in vivo NR assay cannot be used to accurately estimate reduced-N accumulation but still serves as a useful assay for relative differences in treatment conditions.

/pdf/nitrate-reduction-by-roots-of-soybean-glycine-max-l-merr-5drvo4khc1.pdf

Steven J. Crafts-Brandner

Papers

Physiological N response of field-grown maize hybrids (Zea mays L.) with divergent yield potential and grain protein concentration.

Differential Senescence of Maize Hybrids following Ear Removal : I. Whole Plant.

Differential Senescence of Maize Hybrids following Ear Removal II. Selected Leaf

Effects of Pod Removal on Metabolism and Senescence of Nodulating and Nonnodulating Soybean Isolines: II. Enzymes and Chlorophyll.

Nitrate Reduction by Roots of Soybean (Glycine max [L.] Merr.) Seedlings.