Increased cytokinin levels in transgenic PSAG12–IPT tobacco plants have large direct and indirect effects on leaf senescence, photosynthesis and N partitioning

We studied the impact of delayed leaf senescence on the functioning of plants growing under conditions of nitrogen remobilization. Interactions between cytokinin metabolism, Rubisco and protein levels, photosynthesis and plant nitrogen partitioning were studied in transgenic tobacco (Nicotiana tabacum L.) plants showing delayed leaf senescence through a novel type of enhanced cytokinin syn‐thesis, i.e. targeted to senescing leaves and negatively auto‐regulated (P_SAG12–IPT), thus preventing developmental abnormalities. Plants were grown with growth‐limiting nitrogen supply. Compared to the wild‐type, endogenous levels of free zeatin (Z)‐ and Z riboside (ZR)‐type cytokinins were increased up to 15‐fold (total ZR up to 100‐fold) in senescing leaves, and twofold in younger leaves of P_SAG12–IPT. In these plants, the senescence‐associated declines in N, protein and Rubisco levels and photosynthesis rates were delayed. Senescing leaves accumulated more (¹⁵N‐labelled) N than younger leaves, associated with reduced shoot N accumulation (–60%) and a partially inverted canopy N profile in P_SAG12–IPT plants. While root N accumulation was not affected, N translocation to non‐senescing leaves was progressively reduced. We discuss potential consequences of these modified sink–source relations, associated with delayed leaf senescence, for plant productivity and the efficiency of utilization of light and minerals.     

Limitations on photosynthesis of competing individuals in stands and the consequences for canopy structure

The canopy structure of a stand of vegetation is determined by the growth patterns of the individual plants within the stand and the competitive interactions among them. We analyzed the carbon gain of individuals in two dense monospecific stands of Xanthium canadense and evaluated the consequences for intra-specific competition and whole-stand canopy structure. The stands differed in productivity, and this was associated with differences in nitrogen availability. Canopy structure, aboveground mass, and nitrogen contents per unit leaf area (N_area) were determined for individuals, and leaf photosynthesis was measured as a function of N_area. These data were used to calculate the daily carbon gain of individuals. Within stands, photosynthesis per unit aboveground mass (P_mass) of individual plants increased with plant height, despite the lower leaf area ratios of taller plants. The differences in P_mass between the tallest most dominant and shortest most subordinate plants were greater in the high-nitrogen than in the low-nitrogen stand. This indicated that competition was asymmetric and that this asymmetry increased with nitrogen availability. In the high-nitrogen stand, taller plants had a higher P_mass than shorter ones, because they captured more light per unit mass and because they had higher photosynthesis per unit of absorbed light. Conversely, in the low-nitrogen stand, the differences in P_mass between plants of different heights resulted only from differences in their light capture per unit mass. Sensitivity analyses revealed that an increase in N_area, keeping leaf area of plants constant, increased whole-plant carbon gain for the taller more dominant plants but reduced carbon gain in the shorter more subordinate ones, which implies that the N_area values of shorter plants were greater than the optimal values for maximum photosynthesis. On the other hand, the carbon gain of all individual plants, keeping their total canopy N constant, was positively related to an increase in their individual leaf area. At the same time, however, increasing the leaf area for all plants simultaneously reduced the carbon gain of the whole stand. This result shows that the optimal leaf area index (LAI), which maximizes photosynthesis of a stand, is not evolutionarily stable because at this LAI, any individual can increase its carbon gain by increasing its leaf area.     

Canopy structure and nitrogen distribution in dominant and subordinate plants in a dense stand of Amaranthus dubius L. with a size hierarchy of individuals

The objective was to investigate how nitrogen allocation patterns in plants are affected by their vertical position in the vegetation (i.e. being either dominant or subordinate). A garden experiment was carried out with Amaranthus dubius L., grown from seed, in dense stands in which a size hierarchy of nearly equally aged individuals had developed. A small number of dominant plants had most of their leaf area in the highest layers of the canopy while a larger number of subordinate plants grew in the shade of their dominant neighbours. Canopy structure, vertical patterns of leaf nitrogen distribution and leaf photosynthetic characteristics were determined in both dominant and subordinate plants. The light distribution in the stands was also measured. Average N contents per unit leaf area (total canopy nitrogen divided by the total leaf area) were higher in the dominant than in the subordinate plants and this was explained by the higher average MPA (leaf dry mass per unit area) of the dominant plants. However, when expressed on a weight basis, average N contents (LNC_av; total canopy N divided by the total dry weight of leaves) were higher in the subordinate plants. It is possible that these higher LNC_av values reflect an imbalance between carbon and nitrogen assimilation with N uptake exceeding its metabolic requirement. Leaf N content per unit area decreased more strongly with decreasing relative photon flux density in the dominant than in the subordinate plants showing that this distribution pattern can be different for plants which occupy different positions in the light gradient in the canopy. The amount of N which is reallocated from the oldest to the younger, more illuminated leaves higher up in the vegetation may depend on the sink strength of the younger leaves for nitrogen. In the subordinate plants, constrained photosynthetic activity caused by shading might have reduced the sink intensity of these leaves.     

Development of the Monsi-Saeki theory on canopy structure and function

BACKGROUND AND AIMS: Monsi and Saeki (1953) published the first mathematical model of canopy photosynthesis that was based on the light attenuation within a canopy and a light response of leaf photosynthesis. This paper reviews the evolution and development of their theory. SCOPE: Monsi and Saeki showed that under full light conditions, canopy photosynthesis is maximized at a high leaf area index (LAI, total leaf area per unit ground area) with vertically inclined leaves, while under low light conditions, it is at a low LAI with horizontal leaves. They suggested that actual plants develop a stand structure to maximize canopy photosynthesis. Combination of the Monsi-Saeki model with the cost-benefit hypothesis in resource use led to a new canopy photosynthesis model, where leaf nitrogen distribution and associated photosynthetic capacity were taken into account. The gradient of leaf nitrogen in a canopy was shown to be a direct response to the gradient of light. This response enables plants to use light and nitrogen efficiently, two resources whose supply is limited in the natural environment. CONCLUSION: The canopy photosynthesis model stimulated studies to scale-up from chloroplast biochemistry to canopy carbon gain and to analyse the resource-use strategy of species and individuals growing at different light and nitrogen availabilities. Canopy photosynthesis models are useful to analyse the size structure of populations in plant communities and to predict the structure and function of future terrestrial ecosystems.     

Scaling sun and shade photosynthetic acclimation of Alocasia macrorrhiza to whole-plant performance – I. Carbon balance and allocation at different daily photon flux densities

We determined the carbon allocation patterns and construction costs of Alocasia macrorrhiza plants grown at different photon flux densities (PFD) as well as the whole-plant carbon gain of these plants at different daily PFDs. Growth at high PFD resulted in thicker leaves with a higher leaf mass per unit area, and increased biomass allocation to petioles and roots, as compared to growth at low PFD. Increased allocation to petioles may have been necessary to support the heavier leaves, whereas increased allocation to roots may have been necessary to supply sufficient water for the higher transpiration rates in high PFD. Root biomass was highly correlated with the daily, whole-plant transpiration rate. Tissue construction costs per unit dry mass were unchanged by acclimation, but, since the mass per unit areas of leaves, roots and petioles all increased, construction costs per unit leaf area were much higher for plants grown at high PFD. On a per unit leaf area basis, daily whole-plant carbon gain measured at high daily PFD was higher in high- than in low-PFD-grown plants. However, on a per unit leaf mass basis, low-PFD-grown plants had a daily carbon gain at least as high as that of high-PFD-grown plants at high daily PFD. At low daily PFD, low-PFD-grown plants maintained an advantage over high-PFD-grown plants in terms of carbon gain because of their larger leaf area ratios. Thus, in terms of carbon gain, low-PFD-grown plants performed better than sun plants at low PFD and as well as high-PFD-grown plants at high PFD, despite their lower photosynthetic capacities per unit area. For high-PFD-grown plants, the higher construction costs per unit leaf area resulted in lower leaf area ratios, which counteracted the advantage of higher photosynthetic rates per unit leaf area.     

Evolutionarily stable leaf area production in plant populations

Using an analytical model, it was shown that for a given amount of nitrogen in the canopy of a stand (N(T)), there exists an evolutionarily stable leaf area index (ES-LAI), and therefore an evolutionarily stable average leaf nitrogen content (n(ES)(av);n(ES)(av) =N(T)/ES-LAI), at which no individual plant in the stand can increase its photosynthesis by changing its leaf area. It was also shown that this ES-LAI is always greater than the optimal LAI that maximizes photosynthesis per unit N(T) of the stand. This illustrates that the canopy structure that maximizes photosynthesis of a population is not the same as the canopy structure that maximizes photosynthesis of individuals within a population. It was further derived that the ES-LAI at given N(T) increases with the ratio between the light-saturated photosynthesis and the N content per unit leaf area (leaf-PPNUE) and that it decreases with the canopy extinction coefficient for light (K(L)), the light availability and the apparent quantum yield (phi). These hypotheses were tested by comparing calculated ES-LAI and n(ES)(av) values to actual LAIs and leaf N contents measured for stands of a large variety of herbaceous plants. There was a close correspondence between the calculated and measured values. As predicted by the model, plants with high leaf-PPNUEs produced more leaf area per unit nitrogen than those with low leaf-PPNUEs while plants with horizontal leaves, forming stands with higher K(L) values, produced less leaf area than those with more vertically inclined leaves. These results suggest that maximization of individual plant photosynthesis per unit of nitrogen plays an important role in determining leaf area production of plants and the resulting canopy structure of stands of vegetation. They further suggest this optimization to be a mechanism by which leaf traits such as leaf-PPNUE and leaf inclination angle are causally related to structural characteristics of the population, i.e. the leaf area index of the stand.     

A gene encoding an acyl hydrolase is involved in leaf senescence in Arabidopsis

SAG101, a leaf senescence-associated gene, was cloned from an Arabidopsis leaf senescence enhancer trap line and functionally characterized. Reporter gene and RNA gel blot analyses revealed that SAG101 was not expressed until the onset of senescence in leaves. A recombinant SAG101 fusion protein overexpressed in Escherichia coli displayed acyl hydrolase activity. Antisense RNA interference in transgenic plants delayed the onset of leaf senescence for approximately 4 days. Chemically induced overexpression of SAG101 caused precocious senescence in both attached and detached leaves of transgenic Arabidopsis plants. These data suggest that SAG101 plays a significant role in leaf senescence.     

Carbon gain in a multispecies canopy: the role of specific leaf area and photosynthetic nitrogen-use efficiency in the tragedy of the commons

Models have been formulated for monospecific stands in which canopy photosynthesis is determined by the vertical distribution of leaf area, nitrogen and light. In such stands, resident plants can maximize canopy photosynthesis by distributing their nitrogen parallel to the light gradient, with high contents per unit leaf area at the top of the vegetation and low contents at the bottom. Using principles from game theory, we expanded these models by introducing a second species into the vegetation, with the same vertical distribution of biomass and nitrogen as the resident plants but with the ability to adjust its specific leaf area (SLA, leaf area:leaf mass). The rule of the game is that invaders replace the resident plants if they have a higher plant carbon gain than those of the resident plants. We showed that such invaders induce major changes in the vegetation. By increasing their SLA, invading plants could increase their light interception as well as their photosynthetic nitrogen-use efficiency (PNUE, the rate of photosynthesis per unit organic nitrogen). By comparison with stands in which canopy photosynthesis is maximized, those invaded by species of high SLA have the following characteristics: (1) the leaf area index is higher; (2) the vertical distribution of nitrogen is skewed less; (3) as a result of the supra-optimal leaf area index and the more uniform distribution of nitrogen, total canopy photosynthesis is lower. Thus, in dense canopies we face a classical tragedy of the commons: plants that have a strategy to maximize canopy carbon gain cannot compete with those that maximize their own carbon gain. However, because of this strategy, individual as well as total canopy carbon gain are eventually lower. We showed that it is an evolutionarily stable strategy to increase SLA up to the point where the PNUE of each leaf is maximized.     

Vertical variation in leaf gas exchange parameters for a Southeast Asian tropical rainforest in Peninsular Malaysia

Vertical variation in leaf gas exchange characteristics of trees grown in a lowland dipterocarp forest in Peninsular Malaysia was investigated. Maximum net photosynthetic rate, stomatal conductance, and electron transport rate of leaves at the upper canopy, lower canopy, and forest floor were studied in situ with saturated condition photosynthetic photon flux density. The dark respiration rate of leaves at the various heights was also studied. Relationships among gas exchange characteristics, and also with nitrogen content per unit leaf area and leaf dry matter per area were clearly detected, forming general equations representing the vertical profile of several important parameters related to gas exchange. Numerical analysis revealed that the vertical distribution of gas exchange parameters was well determined showing both larger carbon gain for the whole canopy and at the same time positive carbon gain for the leaves of the lowest layer. For correct estimation of gas exchange at both leaf and canopy scales using multi-layer models, it is essential to consider the vertical distribution of gas exchange parameters with proper scaling coefficients.     

Leaf area index, canopy structure and photosynthesis of red clover (Trifolium pratense L.)

Abstract. The influence of leaf age, total leaf area and its dispersion in space on canopy photosynthesis were studied using microswards of red clover ( Trifolium pratense L.) which were established in the greenhouse. Two varieties, Renova (flowering) and Molstad (non-flowering), were sown in separate plastic boxes at densities of 225, 400 and 625 plants per m².
Vertical distribution of photosynthetically active radiation (PAR), leaf area, leaf age and ¹⁴CO₂-fixation were determined periodically. Net photosynthesis and dark respiration of canopies were measured. Maximum photosynthetic capacity of individual leaves was measured on plants taken from the intact canopy or from plants where shading of the growing leaves had been prevented.
Net photosynthetic rate of canopies increased linearly with leaf area index (LAI) up to an LAI of 3.5 and then declined at higher LAI, independent of variety and sowing density. Below the optimum LAI, net photosynthesis depended mainly on interception of PAR. Decrease in canopy photosynthesis above the optimum LAI was due to a higher proportion of old leaves with decreased photosynthetic capacity, and not to an increase in respiring plant parts. It is concluded that LAI and position of leaf age categories in the canopy are more important than vertical distribution of leaf area in determining canopy photosynthesis of red clover.     

Effects of P(SAG12)-IPT gene expression on development and senescence in transgenic lettuce

An ipt gene under control of the senescence-specific SAG12 promoter from Arabidopsis (P(SAG12)-IPT) significantly delayed developmental and postharvest leaf senescence in mature heads of transgenic lettuce (Lactuca sativa L. cv Evola) homozygous for the transgene. Apart from retardation of leaf senescence, mature, 60-d-old plants exhibited normal morphology with no significant differences in head diameter or fresh weight of leaves and roots. Induction of senescence by nitrogen starvation rapidly reduced total nitrogen, nitrate, and growth of transgenic and azygous (control) plants, but chlorophyll was retained in the lower (outer) leaves of transgenic plants. Harvested P(SAG12)-IPT heads also retained chlorophyll in their lower leaves. During later development (bolting and preflowering) of transgenic plants, the decrease in chlorophyll, total protein, and Rubisco content in leaves was abolished, resulting in a uniform distribution of these components throughout the plants. Homozygous P(SAG12)-IPT lettuce plants showed a slight delay in bolting (4-6 d), a severe delay in flowering (4-8 weeks), and premature senescence of their upper leaves. These changes correlated with significantly elevated concentrations of cytokinin and hexoses in the upper leaves of transgenic plants during later stages of development, implicating a relationship between cytokinin and hexose concentrations in senescence.     

Leaf senescence is delayed in tobacco plants expressing the maize homeobox gene knotted1 under the control of a senescence-activated promoter.

Leaf senescence is an active process involving remobilization of nutrients from senescing leaves to other parts of the plant. Whereas senescence is accompanied by a decline in leaf cytokinin content, supplemental cytokinin delays senescence. Plants that overexpress isopentenyl transferase (ipt), a cytokinin-producing gene, or knotted1 (kn1), a homeobox gene, have many phenotypes in common. Many of these phenotypes are characteristic of altered cytokinin physiology. The effect of kn1 on leaf senescence was tested by driving its expression using the promoter of the senescence-associated gene SAG12. SAG:kn1 tobacco plants showed a marked delay in leaf senescence but otherwise developed normally. The delay in senescence was revealed by an increase in chlorophyll content in SAG:kn1 leaves relative to leaves of the control plants and by a decrease in the number of dead leaves. Senescence was also delayed in detached leaves of SAG:kn1 plants. Delayed senescence was accompanied by increased leaf cytokinin content in older leaves expressing kn1. These experiments extend the current understanding of kn1 function and suggest that in addition to mediating meristem maintenance, kn1 is capable of regulating the onset of senescence in leaves.     

重庆石灰岩地区主要木本植物叶片性状及养分再吸收特征

以重庆石灰岩地区15种常绿木本植物和14种落叶木本植物为研究对象,对两种生活型植物叶片衰老前后叶干物质含量(LDMC)、比叶面积(SLA)和叶片厚度(LT)进行了比较,并采用不同的计算方法(单位质量叶片养分含量、单位面积叶片养分含量)分析了两类植物叶片衰老前后养分含量及再吸收特征,最后对养分再吸收效率与其他叶性状因子之间的关系进行了相关分析。结果表明:常绿植物成熟叶LDMC、LT及衰老叶LT显著低于落叶植物,落叶植物成熟叶和衰老叶SLA均显著高于常绿植物(P0.05);基于单位质量叶片计算的养分含量,常绿植物成熟和衰老叶N、P量均低于落叶植物,而基于单位面积叶片计算的N、P含量则表现出相反的趋势;基于不同方法计算的N、P再吸收效率差异不明显,其中常绿植物基于单位质量叶片养分含量计算的N、P平均再吸收效率为39.42%、43.79%,落叶植物的为24.08%、33.59%;常绿和落叶植物N、P再吸收效率与LDMC、SLA、LT和成熟叶N、P含量之间没有显著相关性,但与衰老叶养分含量存在显著负相关(P0.05)。研究发现,无论是常绿植物还是落叶植物,衰老叶N、P含量均较低,表明石灰岩地区植物具有较高的养分再吸收程度。     

Expression of the beta-oxidation gene 3-ketoacyl-CoA thiolase 2 (KAT2) is required for the timely onset of natural and dark-induced leaf senescence in Arabidopsis

The onset of leaf senescence is regulated by a complex mechanism involving positive and negative regulators. Among positive regulators, jasmonic acid (JA) accumulates in senescing leaves and the JA-insensitive coi1-1 mutant displays delayed leaf senescence in Arabidopsis. A strong activated expression of the gene coding for the JA-biosynthetic beta-oxidation enzyme 3-ketoacyl-CoA thiolase 2 (KAT2) in natural and dark-induced senescing leaves of Arabidopsis thaliana is reported here. By using KAT2::GUS and KAT2::LUC transgenic plants, it was observed that dark-induced KAT2 activation occurred both in excised leaves as well as in whole darkened plants. The KAT2 activation associated with dark-induced senescence occurred soon after a move to darkness, and it preceded the detection of symptoms and the expression of senescence-associated gene (SAG) markers. Transgenic plants with reduced expression of the KAT2 gene showed a significant delayed senescence both in natural and dark-induced processes. The rapid induction of the KAT2 gene in senescence-promoting conditions as well as the delayed senescence phenotype and the reduced SAG expression in KAT2 antisense transgenic plants, point to KAT2 as an essential component for the timely onset of leaf senescence in Arabidopsis.     

Senescence-induced ectopic expression of the A. tumefaciens ipt gene in wheat delays leaf senescence, increases cytokinin content, nitrate influx, and nitrate reductase activity, but does not affect grain yield

The manipulation of cytokinin levels by senescence-regulated expression of the Agrobacterium tumefaciens ipt gene through its control by the Arabidopsis SAG12 (senescence-associated gene 12) promoter is an efficient tool for the prolongation of leaf photosynthetic activity which potentially can affect plant productivity. In the present study, the efficiency of this approach was tested on wheat (Triticum aestivum L.)-a monocarpic plant characterized by a fast switch from vegetative to reproductive growth, and rapid translocation of metabolites from leaves to developing grains after anthesis. When compared with the wild-type (WT) control plants, the SAG12::ipt wheat plants exhibited delayed chlorophyll degradation only when grown under limited nitrogen (N) supply. Ten days after anthesis the content of chlorophyll and bioactive cytokinins of the first (flag) leaf of the transgenic plants was 32% and 65% higher, respectively, than that of the control. There was a progressive increase in nitrate influx and nitrate reductase activity. However, the SAG12::ipt and the WT plants did not show differences in yield-related parameters including number of grains and grain weight. These results suggest that the delay of leaf senescence in wheat also delays the translocation of metabolites from leaves to developing grains, as indicated by higher accumulation of ((15)N-labelled) N in spikes of control compared with transgenic plants prior to anthesis. This delay interferes with the wheat reproductive strategy that is based on a fast programmed translocation of metabolites from the senescing leaves to the reproductive sinks shortly after anthesis.     

A model of dynamics of leaves and nitrogen in a plant canopy: an integration of canopy photosynthesis,leaf life span,and nitrogen use efficiency

A model of dynamics of leaves and nitrogen is developed to predict the effect of environmental and ecophysiological factors on the structure and photosynthesis of a plant canopy. In the model, leaf area in the canopy increases by the production of new leaves, which is proportional to the canopy photosynthetic rate, with canopy nitrogen increasing with uptake of nitrogen from soil. Then the optimal leaf area index (LAI; leaf area per ground area) that maximizes canopy photosynthesis is calculated. If leaf area is produced in excess, old leaves are eliminated with their nitrogen as dead leaves. Consequently, a new canopy having an optimal LAI and an optimal amount of nitrogen is obtained. Repeating these processes gives canopy growth. The model provides predictions of optimal LAI, canopy photosynthetic rates, leaf life span, nitrogen use efficiency, and also the responses of these factors to changes in nitrogen and light availability. Canopies are predicted to have a larger LAI and a higher canopy photosynthetic rate at a steady state under higher nutrient and/or light availabilities. Effects of species characteristics, such as photosynthetic nitrogen use efficiency and leaf mass per area, are also evaluated. The model predicts many empirically observed patterns for ecophysiological traits across species.     

Implications of shoot structure on the rate of photosynthesis at different levels in a coniferous canopy using a model incorporating grouping and penumbra

1. The connection between high leaf area index (LAI) and photosynthetic production with two attributes of coniferous canopy structure: small leaf size and grouping of needles on shoots, was analysed using a simulation model.
2. The small size of conifer needles gives rise to penumbras, which even out the distribution of direct sunlight on the leaf area and thereby act to increase the rate of canopy photosynthesis per unit of LAI.
3. Grouping, by producing a non-uniform distribution of leaf area, causes a decrease in total canopy light interception at any given LAI, but improves the photosynthetic light capture by shoots in the lower canopy.
4. Application of the model on a case study showed that: (a) grouping had a negative effect on the rate of photosynthesis in the upper canopy, but deeper down in the canopy the situation was reversed; (b) in the lower canopy, photosynthetic rates were up to 10 times higher as a result from the combined effect of grouping and penumbra; (c) grouping did not improve the rate of canopy photosynthesis per unit of LAI, however, it can have a positive effect on the total photosynthetic production by allowing a higher productive LAI to be maintained; (d) penumbra, on the other hand, increased the rate of canopy photosynthesis by as much as 40% for moderate values of the LAI.     

Mean residence time of leaf number, area, mass, and nitrogen in canopy photosynthesis

Mean residence time (MRT) of plant nitrogen (N), which is an indicator of the expected length of time N newly taken up is retained before being lost, is an important component in plant nitrogen use. Here we extend the concept MRT to cover such variables as leaf number, leaf area, leaf dry mass, and nitrogen in the canopy. MRT was calculated from leaf duration (i.e., time integral of standing amount) divided by the total production of leaf variables. We determined MRT in a Xanthium canadense stand established with high or low N availability. The MRT of leaf number may imply longevity of leaves in the canopy. We found that the MRT of leaf area and dry mass were shorter than that of leaf number, while the MRT of leaf N was longer. The relatively longer MRT of leaf N was due to N resorption before leaf shedding. The MRT of all variables was longer at low N availability. Leaf productivity is the rate of canopy photosynthesis per unit amount of leaf variables, and multiplication of leaf productivity by MRT gives the leaf photosynthetic efficiency (canopy photosynthesis per unit production of leaf variables). The photosynthetic efficiency of leaf number implies the lifetime carbon gain of a leaf in the canopy. The analysis of plant-level N use efficiency by evaluating the N productivity and MRT is a well-established approach. Extension of these concepts to leaf number, area, mass, and N in the canopy will clarify the underlying logic in the study of leaf life span, leaf area development, and dry mass and N use in canopy photosynthesis.     

Leaf senescence and N uptake parameters as selection traits for nitrogen efficiency of oilseed rape cultivars

The cultivation of N-efficient oilseed rape cultivars could contribute to a reduction of the large N balance surpluses of this crop. To facilitate the breeding process of N-efficient cultivars, the identification of secondary plant traits correlating with N efficiency is necessary. The objectives of this study were to investigate leaf senescence and N uptake parameters of oilseed rape cultivars and doubled haploid (DH) lines with contrasting N efficiency in a short-term nutrient solution experiment and to relate these results to their performance in field experiments. In the nutrient solution experiment, genotypes differed in leaf senescence of fully expanded leaves and maximum N uptake rate per unit root length under low N supply. A high maximum N uptake rate seemed to have contributed to delayed leaf senescence by enhancing N accumulation in leaves. Also in the field experiments, genotypes differed in leaf senescence after flowering at limiting N supply. Additionally, the most N-efficient DH line was able to adapt leaf photosynthetic capacity to the low-light conditions in the canopy during flowering. N efficiency (grain yield at limiting N supply) was positively correlated with delayed leaf senescence both in nutrient solution and field experiments. It is concluded that important leaf and root traits of N-efficient cultivars are expressed in short-term nutrient solution experiments, which may facilitate the selection of N-efficient cultivars.     

Canopy dynamics and carbon gain in response to soil water availability in Encelia frutescens gray,a drought-deciduous shrub

Summary The production and longevity of leaves of Encelia frutescens Gray, a drought-deciduous subshrub of the Mohave and Sonoran Deserts, were followed during the summer and fall of 1983 in an experimental field garden. The relationships between seasonally changing plant water status, extent of canopy development, and photosynthetic capacity per unit leaf area were determined. Maximum leaf life spans during a summer activity period were between 3 and 4 months, with the great majority living between 1 and 3 months. Leaf production occurred synchronously in well defined cohorts triggered by precipitation events. Extensive leaf turnover occurred during the summer period even though the plants remained in continuous leaf. Turnover was most pronounced when precipitation triggered the production of new leaf cohorts.Five weeks were required for plants to reach maximum canopy development when renewed soil-water availability followed a prolonged drought. Photosynthetic capacity per unit leaf area recovered much sooner than total leaf area, and submaximal leaf area development was the major factor limiting whole-plant carbon gain during a leaf-flushing period lasting several weeks. As the soil began to dry out, physiological capacity declined more rapidly than leaf area, and became the primary limiting factor to whole plant carbon gain.