Hypobaric Control of Ethylene-Induced Leaf Senescence in Intact Plants of Phaseolus vulgaris L

A controlled atmospheric-environment system (CAES) designed to sustain normal or hypobaric ambient growing conditions was developed, described, and evaluated for its effectiveness as a research tool capable of controlling ethylene-induced leaf senescence in intact plants of Phaseolus vulgaris L.

Senescence was prematurely-induced in primary leaves by treatment with 30 parts per million ethephon. Ethephon-derived endogenous ethylene reached peak levels within 6 hours at 26°C. Total endogenous ethylene levels then temporarily stabilized at approximately 1.75 microliters per liter from 6 to 24 hours. Thereafter, a progressive rise in ethylene resulted from leaf tissue metabolism and release. Throughout the study, the endogenous ethylene content of ethephon-treated leaves was greater than that of nontreated leaves.

Subjecting ethephon-treated leaves to atmospheres of 200 millibars, with O₂ and CO₂ compositions set to approximate normal atmospheric partial pressures, prevented chlorophyll loss. Alternately, subjecting ethephon-treated plants to 200 millibars of air only partially prevented chlorophyll loss. Hypobaric conditions (200 millibars), with O₂ and CO₂ at normal atmospheric availability, could be delayed until 48 hours after ethephon treatment and still prevent most leaf senescence. In conclusion, hypobaric conditions established and maintained within the CAES prevented ethylene-induced senescence (chlorosis) in intact plants, provided O₂ and CO₂ partial pressures were maintained at levels approximating normal ambient availability.

An unexpected increase in endogenous ethylene was detected within nontreated control leaves 48 hours subsequent to relocation from winter greenhouse conditions (latitude, 42°00″ N) to the CAES operating at normal ambient pressure. The longer photoperiod and/or higher temperature utilized within the CAES are hypothesized to influence ethylene metabolism directly and growth-promotive processes (e.g. response thresholds) indirectly.

     

Whole Plant and Leaf Steady State Gas Exchange during Ethylene Exposure in Xanthium strumarium L

The effects of ethylene evolved from ethephon on leaf and whole plant photosynthesis in Xanthium strumarium L. were examined. Ethylene-induced epinasty reduced light interception by the leaves of ethephon treated plants by up to 60%. Gas exchange values of individual, attached leaves under identical assay conditions were not inhibited even after 36 hours of ethylene exposure, although treated leaves required a longer induction period to achieve steady state photosynthesis. The speed of translocation of recently fixed ¹¹C-assimilate movement was not seriously impaired following ethephon treatment; however, a greater proportion of the assimilate was partitioned downward toward the roots. Within 24 hours of ethephon treatment, the whole plant net carbon exchange rate expressed on a per plant basis or a leaf area basis had dropped by 35%. The apparent inhibition of net carbon exchange rate was reversed by physically repositioning the leaves with respect to the light source. Ethylene exposure also inhibited expansion of young leaves which was partially reversed when the leaves were repositioned. The data indicated that ethylene indirectly affected net C gain and plant growth through modification of light interception and altered sink demand without directly inhibiting leaf photosynthesis.     

Role of aminocyclopropane-l-carboxylic acid in ethylene release by distal tissues following localized application of ethephon in Cucurbita pepo

The application of ethephon to a single leaf of Cucurbita pepo L. cv. Trailing Marrow plants caused a huge increase in ethylene production from the treated organ and an increased rate of ethylene production from other parts of the plant. These increases were particularly marked in the shoot apex and expanding leaf. Prior treatment with aminoethoxyvinylglycine (AVG), an ethylene biosynthesis inhibitor, blocked the increased production of ethylene at sites distant from the point of ethephon application. This strongly suggests that the increased ethylene production at these distant sites is due to ethylene biosynthesis and not a result of the translocation of ethylene released by the breakdown of ethephon at the site of application. Assays of 1-aminocyclopropane-l-carboxylic acid (ACC), an ethylene precursor, showed that it increased substantially after ethephon application but was at undetectable levels in the presence of AVG. It is proposed that the application of ethephon stimulates ethylene biosynthesis, but that transport through the plants is effected by ACC which is then converted to ethylene at the shoot apex and leaves.     

Carbon Partitioning in Mature Leaves of Pepper: Effects of Daylength

Grange, R. 1. 1985. Carbon partitioning in mature leaves ofpepper: effects of daylength.—J. exp. Bot. 36: 1749–1759. The partitioning of recently fixed carbon has been examinedin mature pepper leaves grown in 6, 10 or 14 h photoperiodsat different irradiances chosen to give similar radiation integralsand in a 6 h photoperiod at the lowest of these irradiances.The partitioning of carbon into export, starch, sugars and respirationwas followed over the photopenod and the subsequent night ina mature leaf. The maximum export rate during the day (approximately 18 µgC cm^–2 leaf h^–1) was not significantly differentamong the treatments. Net photosynthesis rate was directly relatedto irradiance; the proportion of net photosynthesis exportedduring the day was 33% in 6-h days and 57% in 14-h days. Leafstarch accumulation (as a proportion of net photosynthesis rate)increased slightly when plants were grown in 6-h days. The remobilization of starch and sugars at night allowed exportrates to remain similar over 24 h when plants were grown in10-h or 14-h photoperiods. Leaves grown in 6-h days showed nosignificant changes in export rate during the first few hoursof night but exhausted their starch reserves during the nightand export rates declined. Sucrose and hexose levels decreased at the onset of darkness,but did not fall below 40 µg cm^–2 in plants grownin 10-h or 14-h photoperiods; when this level was reached after3–4 h of darkness, starch breakdown began. In leaves grownin both 6-h treatments, sucrose levels fell below 40 µgcm^–2 when starch reserves were depleted during the nightand the export rate decreased concurrently. The results are discussed in relation to the control of exportand starch metabolism in the leaf. Key words: Pepper, partitioning, daylength     

The role of sink demand in carbon partitioning and photosynthetic reinvigoration following shoot decapitation

Photosynthesis, growth, and carbon partitioning of vigorous coppice shoots were compared with the slower growing intact shoots of Populus maximowiczii × nigra L. MN9 to determine the relationship between carbon partitioning and photosynthetic rate. Relative height growth rate of coppice shoots was 2.2 times that of intact shoots with net photosynthetic rate 1.9 times that of intact shoots. Coppice leaves exported a larger proportion of newly-fixed assimilate (11% compared with 6%) after a 4-h chase. The greater export from coppice leaves was correlated with a greater proportion of [¹⁴C]-labelled photosynthate deposited as starch in stems 4 cm below the point of label application. Coppice leaf assimilate levels were reduced to 15% that of leaves on intact plants, but coppice leaves had twice the concentration of labelled sucrose. Carbohydrates constituted 55% of the water-soluble [¹⁴C]-labelled photosynthate in leaves of coppice shoots compared with 40% in intact shoots. The results suggest that carbon allocation and partitioning in coppice shoots were altered towards production and export of new assimilate, and support the hypothesis that photosynthetic rate is responsive to sink demand for assimilates.     

Partitioning of 14C Labelled Assimilates in Arctium tomentosum

Plants of the biennial Arctium tomentosum were grown from seedto seed-set in an open field under three different treatments:control plants receiving full light intensity, plants with aleaf area reduced by 45 per cent, and shaded plants receivingonly 20 per cent of natural illumination. At various stagesof development the youngest fully expanded leaf of one plantin each treatment was exposed to ¹⁴CO₂ for half an hour. Subsequentdistribution of labelled assimilates in various plant partswas determined after eight hours. In the first year, the mostdominant sink was the tap root irrespective of variation inassimilate supply. During the production of new vegetative growthin the second season, a larger amount of radioactive photosynthatewas recovered from above ground parts, especially during formationof lateral branches. Seed filling consumed 80–90 per centof labelled carbon exported from the exposed leaf. In the secondyear, the most pronounced difference between treatments wasin the degree of apical dominance, being highest in shaded plantsand lowest in the plants with cut leaves. Results from ¹⁴C experimentsagreed fairly well with a ‘partitioning coefficient’derived from a growth analysis of plants grown independentlyunder the same experimental conditions. Reasons for discrepanciesbetween the ¹⁴C results and the partitioning coefficient arediscussed. Arctium tomentosum, burdock, variation in assimilate supply, assimilate distribution, ¹⁴CO₂, labelling, growth analysis     

NADPH oxidase inhibitor diphenyleneiodonium and reduced glutathione mitigate ethephon-mediated leaf senescence,H2O2 elevation and senescence-associated gene expression in sweet potato (Ipomoea batatas)

Ethephon, an ethylene releasing compound, promoted leaf senescence, H₂O₂ elevation, and senescence-associated gene expression in sweet potato. It also affected the glutathione and ascorbate levels, which in turn perturbed H₂O₂ homeostasis. The decrease of reduced glutathione and the accumulation of dehydroascorbate correlated with leaf senescence and H₂O₂ elevation at 72 h in ethephon-treated leaves. Exogenous application of reduced glutathione caused quicker and significant increase of its intracellular level and resulted in the attenuation of leaf senescence and H₂O₂ elevation. A small H₂O₂ peak produced within the first 4 h after ethephon application was also eliminated by reduced glutathione. Diphenyleneiodonium (DPI), an NADPH oxidase inhibitor, delayed leaf senescence and H₂O₂ elevation at 72 h, and its influence was effective only within the first 4 h after ethephon treatment. Ethephon-induced senescence-associated gene expression was repressed by DPI and reduced glutathione at 72 h in pretreated leaves. Leaves treated with l-buthionine sulfoximine, an endogenous glutathione synthetase inhibitor, did enhance senescence-associated gene expression, and the activation was strongly repressed by reduced glutathione. In conclusion, ethephon-mediated leaf senescence, H₂O₂ elevation and senescence-associated gene expression are all alleviated by reduced glutathione and NADPH oxidase inhibitor DPI. The speed and the amount of intracellular reduced glutathione accumulation influence its effectiveness of protection against ethephon-mediated effects. Reactive oxygen species generated from NADPH oxidase likely serves as an oxidative stress signal and participates in ethephon signaling. The possible roles of NADPH oxidase and reduced glutathione in the regulation of oxidative stress signal in ethephon are discussed.     

双台河口翅碱蓬生长与根、茎、叶碳、氮的分配

生物个体生长过程是其同化作用和异化作用过程中物质和能量吸收、转运、储存、排出平衡的综合结果,体现了生物对其生存环境的适应。对采集于双台河口潮滩湿地的翅碱蓬(Suaeda heteroptera)进行株高和根、茎、叶碳、氮含量测定的结果显示,以AIC_C和adj-R~2为准则选择的翅碱蓬株高生长模拟方程为Logistic方程。采用多模型推断确定的双台河口翅碱蓬平均理论渐近株高H_∞为(38.11±2.59)cm。翅碱蓬根、茎、叶碳含量分别为(37.92±4.40)%、(39.98±3.12)%、(28.27±3.41)%;氮含量分别为(0.68±0.35)%、(0.94±0.31)%、(1.26±0.19)%。仅根碳、氮含量与株高间呈显著负相关生长关系。翅碱蓬根碳、氮相对含量分别为97.8±2.1、90.2±9.2,与株高(8.1—36.6 cm)相比,翅碱蓬根碳、氮积累可能受到了环境条件限制。     

The Influence of Leaf Position and Defoliation on the Assimilation and Translocation of Carbon in White Clover (Trifolium repens L.) 1. Carbon Distribution Patterns

The assimilation of carbon (C) by, and distribution of ¹⁴C from,leaves at each end of an unbroken sequence of ten mature leaveson the main stolon of clonal plants of white clover (Trifoliumrepens L.) were measured to identify intra-plant factors determiningthe direction of C movement from leaves. Leaves at two intermediatepositions were also measured. Localized movement of ¹⁴C to sinks at the same node as, or atthe one to two nodes immediately behind, the fed leaf accountedfor 40–50% of the total ¹⁴C exported by all measured leaves.A further 50–60% of exported ¹⁴C was therefore availablefor more-distant sinks, and the direction of translocation ofthis C was determined by the relative total strength or demand(number x size x rate of activity or growth) of sinks forwardof, or behind, the leaf in question. Thus 85% of the ¹⁴C exportedfrom the youngest measured leaf moved toward the base of thestolon, while about 60% of the ¹⁴C exported from the oldestleaf moved acropetally. Defoliating plants to leave just one mature leaf on the mainstolon (at any one of the same four positions studied in undefoliatedplants), and no leaves on branches, resulted in: (1) increasednet photosynthetic rate in all residual leaves: (2) increased%export of fixed C from one of the four leaves; (3) increasedexport to the main stolon apex from all except the eldest leaf;(4) increased export to branches from three of the four leaves;and (5) decreased export to stolon tissue and roots from allleaves, within 3 d of defoliation. These responses would seemto ensure the fastest possible replacement of lost leaf areaand, thus, restoration of homeostatic growth. The observed patternsof C assimilation and distribution in both undefoliated anddefoliated white clover plants are consistent with the generalrules of source-sink theory; the distance between sources andcompeting sinks, and relative sink strength, emerge as the mostimportant intra-plant factors governing C movement. These resultsemphasize the need to consider plant morphology, and the modularnature of plant growth, when interpreting patterns of resourceallocation in clonal plants, or plant responses to stressessuch as partial defoliation. Trifolium repens L, white clover, photosynthesis, assimilate translocation, defoliation     

Partitioning of 14C-labeled photosynthate to allelochemicals and primary metabolites in source and sink leaves of aspen: evidence for secondary metabolite turnover

Theories on allelochemical concentrations in plants are often based upon the relative carbon costs and benefits of multiple metabolic fractions. Tests of these theories often rely on measuring metabolite concentrations, but frequently overlook priorities in carbon partitioning. We conducted a pulse-labeling experiment to follow the partitioning of ¹⁴CO₂-labeled photosynthate into ten metabolic pools representing growth and maintenance (amino acids, organic acids, lipids plus pigments, protein, residue), defense (phenolic glycosides, methanol:water and acetone-soluble tannins/phenolics), and transport and storage (sugars and starch) in source and importing sink leaves of quaking aspen (Populus tremuloides). The peak period of ¹⁴C incorporation into sink leaves occurred at 24 h. Within 48 h of labeling, the specific radioactivity (dpm/mg dry leaf weight) of phenolic glycosides declined by over one-third in source and sink leaves. In addition, the specific radioactivity in the tannin/phenolic fraction decreased by 53% and 28% in source and sink leaves, respectively. On a percent recovery basis, sink leaves partitioned 1.7 times as much labeled photosynthate into phenolic glycosides as source leaves at peak ¹⁴C incorporation. In contrast, source leaves partitioned 1.8 times as much ¹⁴C-labeled photosynthate into tannins/phenolics as importing sink leaves. At the end of the 7-day chase period, sink leaves retained 18%, 52%, and 30% of imported ¹⁴C photosynthate, and labeled source leaves retained 15%, 66%, and 19% of in situ photosynthate in metabolic fractions representing transport and storage, growth and maintenance, and defense, respectively. Analyses of the phenolic fractions showed that total phenolics were twice as great and condensed tannins were 1.7 times greater in sink than in source leaves. The concentration of total phenolics and condensed tannins did not change in source and sink leaves during the 7-day chase period. Received: 31 July 1998 / Accepted: 8 February 1999     

Polarity of the stem and ontogenetic changes in sunflow (Helianthus annuus L.) leaves in relation to endogenou cytokinins and ethylene

The content of endogenous cytokinin-like substances and the release of ethylene were determined in leaves of different insertion of sunflower plants during their ontogeny. The content of cytokinin-like substances was highest in the leaves on the middle part of the stem (that is in leaves just before full expansion), with a decrease occurring both towards the base and the apex of the stem, when followed at four growth phases (vegetative plants, plants with inflorescence diameter up to 0.5 cm, plants with inflorescence diameter up to 3 cm, and plants in flower). Changes in the content of cytokinin-like substances during the ontogeny of the leaf also corresponded to this pattern. Data obtained with the leaf at the third node from the basis of the stem showed that the level of cytokinin-like substances first sharply increased, and then after reaching maximal value (at the time when leaf blade area reached approximately 70 % of the final value) slowly and continuously decreased. The highest amount of ethylene released from the leaves was recorded in basal leaves and then also in apical leaves, whereas the leaves with the largest blade area situated at the central part of the stem released the lowest amount of ethylene. This pattern was repeatedly found at all four selected growth phases of sunflower plants.     

Routes of Ethephon Uptake in Pineapple (Ananas comosus) and Reasons for Failure of Flower Induction

Ethylene-releasing agents such as ethephon (2-chloroethylphosphonic acid) are used widely to induce flowering in pineapples (Ananas comosus (L.) Merrill). However, ethephon treatment is less reliable in summer, particularly if plants are treated on abnormally hot days. [¹⁴C]ethephon was used to follow uptake and translocation in leaf tissues. Up to 30% of the ethephon entered the leaf within 4 h, and up to 60% by 24 h. Uptake was dramatically modified by temperature, relative humidity, solution pH, and the surface on which solution droplets were placed. Entry occurred across the leaf cuticle and probably also by way of stomatal pores, and label was recovered at all depths within the leaf. ¹⁴C label entered more rapidly through the abaxial epidermis than through the adaxial epidermis. Low-volume spray applications to whole plants resulted in rapidly drying droplets mainly on the adaxial, distal epidermis and were rather ineffective at inducing flowering, possibly because little ethephon or ethylene reaches the shoot apex. High-volume sprays may facilitate ethephon entry because solution accumulates in leaf axils and hence remains in prolonged contact with abaxial epidermis of leaf bases close to the shoot apex. When poured into the center of the plant, 20% of a normal commercial ethephon dose induced full flowering even under adverse temperatures. It is suggested that high-volume evening spraying and avoidance of hot days may reduce the incidence of flowering failure. Received March 20, 1998; accepted September 6, 1999     

The Influence of Applied Nitrogen on Export and Partitioning of Current Assimilate by Field-grown Potato Plants

Field-grown potatoes were subjected to N deficiency (no appliedN) or received high levels of N (240 kg N ha^–1) at planting.The effects of these treatments were monitored at five stagesduring growth in terms of the allocation of photosynthate withinthe leaf, and the export and partitioning of carbon to differentsinks. N deficiency significantly raised the starch concentrationin all organs of the plants, particularly in leaves and stems,and as a consequence the total amount of starch in the canopyof the low N plants remained greater than that of the high Nplants until approx. 100 days after planting (DAP). The totalamounts of carbohydrates, protein and amino acids were calculatedfor each treatment and these values were used to derive a balancesheet for major reserves. Net losses of reserves occurred fromthe canopy in both treatments in the period 97–133 DAP,although these were shown to represent < 3 per cent of thetotal gain in tuber dry weight for the season. Partitioning of ¹⁴C assimilates was examined in whole plantsand also in single leaves. Reduced partitioning to the tubers,seen in high N plants throughout their growth, was shown tobe due to decreased percentage export by the leaf and accumulationof exported ¹⁴C by the stems. Partitioning to the tubers inlow N plants increased prior to senescence when 87 per centof the fixed ¹⁴C was exported within 24 h, 80 per cent of thisto the tubers. The equivalent values for the high N plants were77 and 60 per cent respectively. Increased percentage exportcoincided with decreased allocation to starch in the leaf, anda link between these processes is suggested. N also significantlyaltered the allocation of ¹⁴C within the leaf and may have influencedthe degradation of starch in the dark to a greater degree thanits synthesis in the light. The enzymes sucrose phosphate synthase (SPS), and starch synthasewere measured concurrently with partitioning. High N plantsshowed higher rates of activities of each of the enzymes althoughboth enzymes showed a similar pattern of development over theseason, irrespective of N treatment. The data are discussed in the light of conflicting reports concerningthe influence of N on translocation and partitioning. ¹⁴C assimilates, carbohydrates, nitrogen, potato (Solanum tuberosum L.), protein     

The application of ethephon (an ethylene releaser) increases growth, photosynthesis and nitrogen accumulation in mustard (Brassica juncea L.) under high nitrogen levels

Ethephon (2-chloroethyl phosphonic acid), an ethylene-releasing compound, influences growth and photosynthesis of mustard (Brassica juncea L. Czern & Coss.). We show the effect of nitrogen availability on ethylene evolution and how this affects growth, photosynthesis and nitrogen accumulation. Ethylene evolution in the control with low N (100 mg N kg(-1) soil) was two-times higher than with high N (200 mg N kg(-1) soil). The application of 100-400 microl x l(-1) ethephon post-flowering, i.e. 60 days after sowing, on plants receiving low or high N further increased ethylene evolution. Leaf area, relative growth rate (RGR), photosynthesis, leaf nitrate reductase (NR) activity and leaf N reached a maximum with application of 200 microl x l(-1) ethephon and high N. The results suggest that the application of ethephon influences growth, photosynthesis and N accumulation, depending on the amount of nitrogen in the soil.     

Influence of gibberellic acid on <Superscript>14</Superscript>CO<Subscript>2</Subscript> metabolism,growth, and production of alkaloids in <Emphasis Type="Italic">Catharanthus roseus</Emphasis>

Changes in growth parameters, carbon assimilation efficiency, and utilization of ¹⁴CO₂ assimilate into alkaloids in plant parts were investigated at whole plant level by treatment of Catharanthus roseus with gibberellic acid (GA). Application of GA (1 000 g m⁻³) resulted in changes in leaf morphology, increase in stem elongation, leaf and internode length, plant height, and decrease in biomass content. Phenotypic changes were accompanied by decrease in contents of chlorophylls and in photosynthetic capacity. GA application resulted in higher % of total alkaloids accumulated in leaf, stem, and root. GA treatment produced negative phenotypic response in total biomass production but positive response in content of total alkaloids in leaf, stem, and roots. ¹⁴C assimilate partitioning revealed that ¹⁴C distribution in leaf, stem, and root of treated plants was higher than in untreated and variations were observed in contents of metabolites as sugars, amino acids, and organic acids. Capacity to utilize current fixed ¹⁴C derived assimilates for alkaloid production was high in leaves but low in roots of treated plants despite higher content of ¹⁴C metabolites such as sugars, amino acids, and organic acids. In spite of higher availability of metabolites, their utilization into alkaloid production is low in GA-treated roots.     

Transport of [2-14C]jasmonic acid from leaves to roots mimics wound-induced changes in endogenous jasmonic acid pools in Nicotiana sylvestris

Jasmonic acid (JA) is part of a long-distance signal-transduction pathway that effects increases in de-novo nicotine synthesis in the roots of Nicotiana sylvestris Speg et Comes (Solanaceae) after leaf wounding. Elevated nicotine synthesis increases whole-plant nicotine pools and makes plants more resistant to herbivores. Leaf wounding rapidly increases JA pools in damaged leaves, and after a 90-min delay, root JA pools also increase. The systemic response in the roots could result from either: (i) the direct transport of JA from wounded leaves, or (ii) JA synthesis or its release from conjugates in roots in response to a second, systemic signal. We synthesized [2-¹⁴C]JA, and applied it to a single leaf in a quantity (189 μg) known to elicit both a whole-plant nicotine and root JA response equivalent to that found in plants subjected to leaf wounding. We quantified radioactive material in JA, and in metabolites both more and less polar than JA, from treated and untreated leaves and roots of plants in eight harvests after JA application. [2-¹⁴C]Jasmonic acid was transported from treated leaves to roots at rates and in quantities equivalent to the wound-induced changes in endogenous JA pools. The [2-¹⁴C]JA that had been transported to the roots declined at the same rate as endogenous JA pools in the roots of plants after leaf wounding. Most of the labeled material applied to leaves was metabolized or otherwise immobilized at the application site, and the levels of [2-¹⁴C]JA in untreated leaves did not increase over time. We measured the free JA pools before and after four different hydrolytic extractions of root and shoot tissues to estimate the size of the potential JA conjugate pools, and found them to be 10% or less of the free JA pool. We conclude that the direct transport of wound-induced JA from leaves to roots can account for the systemic increase in root JA pools after leaf wounding, and that metabolism into less polar structures determines the duration of this systemic increase. However, the conclusive falsification of this hypothesis will require the suppression of all other signalling pathways which could have shoot-to-root transport kinetics similar to that of endogenous JA. Received: 14 April 1997 / Accepted: 9 June 1997     

Influence of Ethephon on Nitrate Reductase, Protein Amino Nitrogen and Nitrate Content of Solanum tuberosum L.

Nitrate reductase activity was stimulated in roots and stems,but suppressed in leaves of potato plants grown in nutrientculture by 30 mg.liter^–1 ethephon applied to the culturesolution. In stems, nitrate reductase activity was stimulatedafter 5 h and by 24 h it was more than two fold that of thecontrol. The magnitude of stimulation by ethephon was less inroots compared to stems. Ethephon treatment enhanced ethyleneproduction by roots, stems and leaves but the level of productionwas not significantly different in these organs. The stimulationof nitrate reductase activity was prevented by cycloheximideand cordycepin suggesting the involvement of new protein synthesis. Ethephon enhanced TCA precipitable protein levels in both rootsand stems while that in leaves was not significantly affected.Amino nitrogen content increased in parallel with protein contentin response to ethephon, with roots exhibiting substantial stimulation.Nitrate accumulation in stem tissues was not affected by ethephontreatment but was increased in roots at 24 and 48 h. Leaf NO³content declined with time in both ethephon-treated and controlplants and after 24 h significantly less NO³ accumulated intreated leaves. These results are explained in terms of ethephonstimulated protein synthesis and increase in cellular metabolismand permeability. (Received August 21, 1984; Accepted January 7, 1985)     

Root Development and Source-Sink Relations in Carrot, Daucus carota L: II. EFFECTS OF ROOT PRUNING ON CARBON ASSIMILATION AND THE PARTITIONING OF ASSIMILATES

Physiological responses to root pruning were investigated bycomparing ¹⁴CO₂ fixation rates, the partitioning of ¹⁴C-labelledassimilate, and soluble and insoluble carbohydrate levels inthe leaves of carrot plants following the removal of some ofthe fibrous roots, or fibrous roots and part of the tap root.Root pruning reduced ¹⁴CO₂ fixation by 28–45% but leafspecific activity (¹⁴C assimilation g-¹ leaf fresh weight) wasunchanged. The proportion of total assimilate exported to theroot system increased following root pruning and this was atthe expense of the developing leaves. In younger plants (wherethe tap root received 10% of the assimilate) the supply of ¹⁴Cto the tap root was maintained in spite of root pruning. However,shortening the tap root to 3 cm in older plants (in which 30%of the fixed 14C was normally exported to the developing storageorgan), reduced its sink capacity and resulted in slightly greaterretention of ¹⁴C in the mature leaves. Greater concentrationsof insoluble carbohydrate were found in the mature leaves followingroot pruning but soluble sugar content was unaffected. Onlysmall differences were observed in the distribution of ¹⁴C betweensoluble and insoluble carbohydrate fractions when plants werefed ¹⁴CO₂ several days after the root pruning operations. Thesephysiological responses were mainly associated with the removalof fibrous roots and support the view that the fibrous rootsystem is more important than the developing storage organ inregulating growth in young carrot plants.     

Nonprotein amino acid composition of flatpea (Lathyrus sylvestris L.) as affected by ethephon seed treatments and seedling fertilization

Use of flatpea (Lathyrus sylvestris L.) as a forage is limited because of nonuniform seed germination and the potentially toxic effects of 2,4-diaminobutyric acid (A₂bu), a nonprotein amino acid found in seeds and vegetative tissues. The effects of ethephon (2-chloroethyl phosphonic acid) on seed germination, amino acid leachates of seeds, and amino acid composition (particularly A₂bu) of seedlings were investigated. Germination of flatpea seeds, imbibed for 16 h in 0, 100, 200, 400, 800, and 1600 mg/L ethephon, did not differ, but amino acid leachates tended to increase up to 200 mg/L ethephon and then decline at higher concentrations. The major amino acid constituents in leachates were A₂bu, 4-aminobutyric acid (Abu), and homoserine (Hse). Dry matter accumulation of seedlings grown from ethephon-treated seeds was reduced for second cuttings grown from ethephon-treated seeds and high nitrogen grown plants. During regrowth, free amino acid accumulation was most pronounced in leaves of plants supplied with high nitrogen. The most abundant free amino acids in flatpea tissues were the same as those in seed leachates, but concentration and relative abundance varied with nitrogen level, plant part, and ethephon treatment. Results suggest that ethephon seed treatments can have persistent effects on the growth and amino acid composition of flatpea seedlings grown under different nitrogen regimes.     

Combined effects of elevated CO2 and soil drought on carbon and nitrogen allocation of the desert shrub Caragana intermedia

Impacts of either elevated CO₂ or drought stress on plant growth have been studied extensively, but interactive effects of these on plant carbon and nitrogen allocation is inadequately understood yet. In this study the response of the dominant desert shrub, Caragana intermedia Kuanget H.c.Fu, to the interaction of elevated CO₂ (700 ± 20 μmol mol⁻¹) and soil drought were determined in two large environmental growth chambers (18 m²). Elevated CO₂ increased the allocation of biomass and carbon into roots and the ratio of carbon to nitrogen (C:N) as well as the leaf soluble sugar content, but decreased the allocation of biomass and carbon into leaves, leaf nitrogen and leaf soluble protein concentrations. Elevated CO₂ significantly decreased the partitioning of nitrogen into leaves, but increased that into roots, especially under soil drought. Elevated CO₂ significantly decreased the carbon isotope discrimination (Δ) in leaves, but increased them in roots, and the ratio of Δ values between root and leaf, indicating an increased allocation into below-ground parts. It is concluded that stimulation of plant growth by CO₂ enrichment may be negated under soil drought, and under the future environment, elevated CO₂ may partially offset the negative effects of enhanced drought by regulating the partitioning of carbon and nitrogen.