水分和养分添加对羊草功能性状和地上生物量的影响

研究水分和养分添加对植物功能性状的影响, 对于揭示植物对环境变化的响应和适应规律至关重要。该文采用盆栽试验的方法, 进行不同水平水分处理(增水50%, 减水50%, 以498 mm降水量作为对照)和养分添加(无养分添加, 单施氮肥, 单施磷肥, 氮磷共施), 研究羊草(Leymus chinensis)的10种功能性状和地上生物量对水分和养分添加的响应。得出以下结论: (1)双因素方差分析结果表明, 水分主效应对羊草株高、分蘖数、茎生物量、叶生物量、叶面积、叶质量、净光合速率、蒸腾速率、水分利用效率存在显著影响; 养分主效应对羊草分蘖数、茎生物量、净光合速率、蒸腾速率、水分利用效率存在显著影响; 水分和养分的交互作用对羊草分蘖数、茎生物量、蒸腾速率、水分利用效率存在显著影响。(2)各功能性状对降水量的响应在不同养分添加水平是不同的, 分蘖数和叶面积在单施氮肥和氮磷共施条件下随降水量增加而增加, 而在无养分添加和单施磷肥条件下无显著变化; 茎生物量在无养分添加、单施氮肥和单施磷肥条件下随降水量增加而增加, 而在氮磷共施条件下无增加趋势; 比叶面积在单施氮肥条件下增水处理显著低于对照组, 而在其他养分添加条件下无明显变化。(3)短期氮磷处理显著影响羊草叶片光合生理性状, 而对叶形态性状影响不显著。(4)羊草地上生物量随降水量的增加呈现上升趋势, 并且在单施氮肥条件下, 增水处理使地上生物量达到最高, 为522.55 g·m ^-2。总之, 羊草的功能性状对降水量增加表现出明显的响应, 响应格局在不同养分条件下不同, 反映了其对水肥环境变化的适应。     

Nitrogen and water addition reduce leaf longevity of steppe species

Background and aims

Changes in supplies of resources will modify plant functional traits. However, few experimental studies have addressed the effects of nitrogen and water variations, either singly or in combination, on functional traits.

Methods

A 2-year field experiment was conducted to test the effects of nitrogen and water addition on leaf longevity and other functional traits of the two dominant (Agropyron cristatum and Stipa krylovii) and three most common species (Cleistogenes squarrosa, Melilotoides ruthenica and Potentilla tanacetifolia) in a temperate steppe in northern China.

Key Results

Additional nitrogen and water increased leaf nitrogen content and net photosynthetic rate, and changed other measured functional traits. Leaf longevity decreased significantly with both nitrogen addition (–6 days in 2007 and –5·4 days in 2008; both P < 0·001) and watering (–13 days in 2007 and –9·9 days in 2008; both P < 0·001), and significant differences in leaf longevity were also found among species. Nitrogen and water interacted to affect leaf longevity and other functional traits. Soil water content explained approx. 70 % of the shifts in leaf longevity. Biomass at both species and community level increased under water and nitrogen addition because of the increase in leaf biomass production per individual plant.

Conclusions

The results suggest that additional nitrogen and water supplies reduce plant leaf longevity. Soil water availability might play a fundamental role in determining leaf longevity and other leaf functional traits, and its effects can be modified by soil nitrogen availability in semi-arid areas. The different responses of species to resource alterations may cause different global change ramifications under future climate change scenarios.     

Interspecific prediction of photosynthetic light response curves using specific leaf mass and leaf nitrogen content: effects of differences in soil fertility and growth irradiance

Background and Aims

Previous work has shown that the entire photosynthetic light response curve, based on both Mitscherlich and Michaelis–Menten functions, could be predicted in an interspecific context through allometric relations linking the parameters of these functions to two static leaf traits: leaf nitrogen (N) content and leaf mass per area (LMA). This paper describes to what extent these allometric relations are robust to changes in soil fertility and the growth irradiance of the plants.

Methods

Plants of 25 herbaceous species were grown under controlled conditions in factorial combinations of low/high soil fertility and low/high growth irradiance. Net photosynthetic rates per unit dry mass were measured at light intensities ranging from 0 to 700 µmol m⁻² s⁻¹ photosynthetically active radiation (PAR).

Key Results

The differing growth environments induced large changes in N, LMA and in each of the parameter estimates of the Mitscherlich and Michaelis–Menten functions. However, the differing growth environments induced only small (although significant) changes in the allometric relationships linking N and LMA to the parameters of the two functions. As a result, 88 % (Mitcherlich) and 89 % (Michaelis–Menten) of the observed net photosynthetic rates over the full range of light intensities (0–700 µmol m⁻² s⁻¹ PAR) and across all four growth environments could be predicted using only N and LMA using the same allometric relations.

Conclusions

These results suggest the possibility of predicting net photosynthetic rates in nature across species over the full range of light intensities using readily available data.     

Mycorrhization and phosphorus nutrition affect water relations and CAM induction by drought in seedlings of Clusia minor

Background and Aims

Crassulacean acid metabolism (CAM) is currently viewed as an adaptation to water deficit. In plants of Clusia minor, which grow mostly on acidic, P-deficient soils, CAM is induced by water deficit. The symbiosis between plants and mycorrhizal fungi alleviates the symptoms of P deficiency and may influence drought resistance. Therefore, the effect of P supply, modified by three different experimental treatments, on the induction of CAM by drought in C. minor was investigated to test the hypothesis that P deficiency will produce greater CAM activity and, in addition, that treatment will modify drought tolerance.

Methods

Seedlings were grown in forest soil sterilized and inoculated with Scutellospora fulgida (SF treatment), sterilized and supplemented with P (Ph treatment) or non-sterilized and containing native mycorrhizae (Nat treatment). Leaf turgor potential (ψ_T) was determined psychrometrically, and CAM activity as nocturnal acid accumulation (ΔH⁺) by titration of dawn and dusk leaf sap.

Key Results

Plant mass and P content were higher in SF and Ph than in Nat seedlings. After 21 d of water deficit, ψ_T increased in SF, decreased in Ph and remained unchanged in Nat, and, after 7 and 14 d of water deficit, ΔH⁺ in Nat was three times higher than at the beginning of drought, whereas in SF and Ph ΔH⁺ was lower than on day 0.

Conclusions

P deficiency in Nat seedlings was ameliorated by inoculation or P addition. The SF and Nat seedlings showed greater tolerance of drought than Ph. P deficiency promoted the induction of CAM by drought in Nat seedlings, whereas P fertilization and mycorrhization did not. Nocturnal acid accumulation was highly and negatively correlated with plant P and N contents, indicating that P and N deficiencies are promoters of CAM in droughted plants of C. minor.Key words: Clusia minor, crassulacean acid metabolism, CAM, mycorrhiza, drought, phosphorus deficiency, nitrogen–water relations     

Optimal use of leaf nitrogen explains seasonal changes in leaf nitrogen content of an understorey evergreen shrub

Background and Aims

Understorey evergreen species commonly have a higher leaf nitrogen content in winter than in summer. Tested here is a hypothesis that such changes in leaf nitrogen content maximize nitrogen-use efficiency, defined as the daily carbon gain per unit nitrogen, under given temperature and irradiance levels.

Methods

The evergreen shrub Aucuba japonica growing naturally at three sites with different irradiance regimes in Japan was studied. Leaf photosynthetic characteristics, Rubisco and leaf nitrogen with measurements of temperature and irradiance monthly at each site were determined. Daily carbon gain was determined as a function of leaf nitrogen content to calculate the optimal leaf nitrogen content that maximized daily nitrogen-use efficiency.

Key Results

As is known, the optimal leaf nitrogen content increased with increasing irradiance. The optimal leaf nitrogen content also increased with decreasing temperature because the photosynthetic capacity per Rubisco decreased. Across sites and months, the optimal leaf nitrogen content was close to the actual leaf nitrogen content and explained the variation in actual leaf nitrogen by 64 %. Sensitivity analysis showed that the effect of temperature on optimal nitrogen content was similar in magnitude to that of irradiance.

Conclusions

Understorey evergreen species regulate leaf nitrogen content so as to maximize nitrogen-use efficiency in daily carbon gain under changing irradiance and temperature conditions.     

Soil microbial properties and plant growth responses to carbon and water addition in a temperate steppe: the importance of nutrient availability

Background

Global climatic change is generally expected to stimulate net primary production, and consequently increase soil carbon (C) input. The enhanced C input together with potentially increased precipitation may affect soil microbial processes and plant growth.

Methodology/Principal Findings

To examine the effects of C and water additions on soil microbial properties and plant growth, we conducted an experiment lasting two years in a temperate steppe of northeastern China. We found that soil C and water additions significantly affected microbial properties and stimulated plant growth. Carbon addition significantly increased soil microbial biomass and activity but had a limited effect on microbial community structure. Water addition significantly increased soil microbial activity in the first year but the response to water decreased in the second year. The water-induced changes of microbial activity could be ascribed to decreased soil nitrogen (N) availability and to the shift in soil microbial community structure. However, no water effect on soil microbial activity was visible under C addition during the two years, likely because C addition alleviated nutrient limitation of soil microbes. In addition, C and water additions interacted to affect plant functional group composition. Water addition significantly increased the ratio of grass to forb biomass in C addition plots but showed only minor effects under ambient C levels. Our results suggest that soil microbial activity and plant growth are limited by nutrient (C and N) and water availability, and highlight the importance of nutrient availability in modulating the responses of soil microbes and plants to potentially increased precipitation in the temperate steppe.

Conclusions/Significance

Increased soil C input and precipitation would show significant effects on soil microbial properties and plant growth in the temperate steppe. These findings will improve our understanding of the responses of soil microbes and plants to the indirect and direct climate change effects.     

Effects of increased nitrogen deposition and precipitation on seed and seedling production of Potentilla tanacetifolia in a temperate steppe ecosystem

Background

The responses of plant seeds and seedlings to changing atmospheric nitrogen (N) deposition and precipitation regimes determine plant population dynamics and community composition under global change.

Methodology/Principal Findings

In a temperate steppe in northern China, seeds of P. tanacetifolia were collected from a field-based experiment with N addition and increased precipitation to measure changes in their traits (production, mass, germination). Seedlings germinated from those seeds were grown in a greenhouse to examine the effects of improved N and water availability in maternal and offspring environments on seedling growth. Maternal N-addition stimulated seed production, but it suppressed seed mass, germination rate and seedling biomass of P. tanacetifolia. Maternal N-addition also enhanced responses of seedlings to N and water addition in the offspring environment. Maternal increased-precipitation stimulated seed production, but it had no effect on seed mass and germination rate. Maternal increased-precipitation enhanced seedling growth when grown under similar conditions, whereas seedling responses to offspring N- and water-addition were suppressed by maternal increased-precipitation. Both offspring N-addition and increased-precipitation stimulated growth of seedlings germinated from seeds collected from the maternal control environment without either N or water addition. Our observations indicate that both maternal and offspring environments can influence seedling growth of P. tanacetifolia with consequent impacts on the future population dynamics of this species in the study area.

Conclusion/Significance

The findings highlight the importance of the maternal effects on seed and seedling production as well as responses of offspring to changing environmental drivers in mechanistic understanding and projecting of plant population dynamics under global change.     

Optimizing nitrogen economy under drought: increased leaf nitrogen is an acclimation to water stress in willow (Salix spp.)

Background and Aims

The major objective was to identify plant traits functionally important for optimization of shoot growth and nitrogen (N) economy under drought. Although increased leaf N content (area basis) has been observed in dry environments and theory predicts increased leaf N to be an acclimation to drought, experimental evidence for the prediction is rare.

Methods

A pedigree of 200 full-sibling hybrid willows was pot-grown in a glasshouse in three replicate blocks and exposed to two water regimes for 3 weeks. Drought conditions were simulated as repeated periods of water shortage. The total leaf mass and area, leaf area efficiency (shoot growth per unit leaf area, E_A), area-based leaf N content (N_A), total leaf N pool (N_L) and leaf N efficiency (shoot growth per unit leaf N, E_N) were assessed.

Key Results

In the water-stress treatment, shoot biomass growth was N limited in the genotypes with low N_L, but increasingly limited by other factors in the genotypes with greatest N_L. The N_A was increased by drought, and drought-induced shift in N_A varied between genotypes (significant G × E). Judged from the E_A–N_A relationship, optimal N_A was 16 % higher in the water-stress compared with the well-watered treatment. Biomass allocation to leaves and shoots varied between treatments, but the treatment response of the leaf : shoot ratio was similar across all genotypes.

Conclusions

It is concluded that N-uptake efficiency and leaf N efficiency are important traits to improve growth under drought. Increased leaf N content (area basis) is an acclimation to optimize N economy under drought. The leaf N content is an interesting trait for breeding of willow bioenergy crops in a climate change future. In contrast, leaf biomass allocation is a less interesting breeding target to improve yield under drought.     

Phenology and growth adjustments of oil palm (Elaeis guineensis) to photoperiod and climate variability

Background and Aims

Oil palm flowering and fruit production show seasonal maxima whose causes are unknown. Drought periods confound these rhythms, making it difficult to analyse or predict dynamics of production. The present work aims to analyse phenological and growth responses of adult oil palms to seasonal and inter-annual climatic variability.

Methods

Two oil palm genotypes planted in a replicated design at two sites in Indonesia underwent monthly observations during 22 months in 2006–2008. Measurements included growth of vegetative and reproductive organs, morphology and phenology. Drought was estimated from climatic water balance (rainfall – potential evapotranspiration) and simulated fraction of transpirable soil water. Production history of the same plants for 2001–2005 was used for inter-annual analyses.

Key Results

Drought was absent at the equatorial Kandista site (0°55′N) but the Batu Mulia site (3°12′S) had a dry season with variable severity. Vegetative growth and leaf appearance rate fluctuated with drought level. Yield of fruit, a function of the number of female inflorescences produced, was negatively correlated with photoperiod at Kandista. Dual annual maxima were observed supporting a recent theory of circadian control. The photoperiod-sensitive phases were estimated at 9 (or 9 + 12 × n) months before bunch maturity for a given phytomer. The main sensitive phase for drought effects was estimated at 29 months before bunch maturity, presumably associated with inflorescence sex determination.

Conclusion

It is assumed that seasonal peaks of flowering in oil palm are controlled even near the equator by photoperiod response within a phytomer. These patterns are confounded with drought effects that affect flowering (yield) with long time-lag. Resulting dynamics are complex, but if the present results are confirmed it will be possible to predict them with models.     

控制降水梯度对荒漠草原优势植物叶功能性状及土壤养分的影响

探究植物叶功能性状随降水梯度的变化规律,对揭示干旱区优势植物对环境变化的响应和适应策略至关重要。以盐池荒漠草原为研究对象,采用遮雨棚和喷灌系统控制降水梯度,分析了优势植物蒙古冰草（Agropyron mongolicum）、短花针茅（Stipa breviflora）及达乌里胡枝子（Lespedeza davurica）叶功能性状变异,以及土壤水分、养分和微生物特性与响应性状间的相关关系。结果表明：HW（增水50%）处理下3个植物LA（叶面积）、LDMC（干物质含量）显著增大,LW处理（减水50%）下短花针茅和达乌里胡枝子LA、LDMC显著减小,降水处理对短花针茅和达乌里胡枝子SLA（比叶面积）影响不显著,LW处理显著提高了蒙古冰草SLA;LW处理显著提高了蒙古冰草和短花针茅LNC（叶氮含量）和LPC（叶磷含量）;HW显著降低了土壤C、N含量,LW和HW均显著减少了真菌数量,而放线菌数量、微生物生物量C、N显著增加;3种优势植物LA均与土壤水分显著正相关,蒙古冰草和短花针茅通过提高SLA、LNC及LNP来适应干旱生境,蒙古冰草和短花针茅LNC及LNP是表征土壤P、微生物生物量有效性的关键指标,达乌里胡枝子通过自我调节养分利用策略来适应C、N、P含量和微生物活性较低的生境,从而决定其在群落中的优势地位。     

Recovery dynamics of growth, photosynthesis and carbohydrate accumulation after de-submergence: a comparison between two wetland plants showing escape and quiescence strategies

Background and Aims

The capacity for fast-growth recovery after de-submergence is important for establishment of riparian species in a water-level-fluctuation zone. Recovery patterns of two wetland plants, Alternanthera philoxeroides and Hemarthria altissima, showing ‘escape’ and ‘quiescence’ responses, respectively, during submergence were investigated.

Methods

Leaf and root growth and photosynthesis were monitored continuously during 10 d of recovery following 20 d of complete submergence. Above- and below-ground dry weights, as well as carbohydrate concentrations, were measured several times during the experiment.

Key Results

Both species remobilized stored carbohydrate during submergence. Although enhanced internode elongation depleted the carbohydrate storage in A. philoxeroides during submergence, this species resumed leaf growth 3 d after de-submergence concomitant with restoration of the maximal photosynthetic capacity. In contrast, some sucrose was conserved in shoots of H. altissima during submergence, which promoted rapid re-growth of leaves 2 d after de-submergence and earlier than the full recovery of photosynthesis. The recovery of root growth was delayed by 1–2 d compared with leaves in both species.

Conclusions

Submergence tolerance of the escape and quiescence strategies entails not only the corresponding regulation of growth, carbohydrate catabolism and energy metabolism during submergence but also co-ordinated recovery of photosynthesis, growth and carbohydrate partitioning following de-submergence.     

土壤水分与遮荫水平对棉花叶片光全特性的影响研究

以盆栽棉花（中棉２３号）做为供试材料,通过人工遮荫,设计了３种遮荫水平（不遮光（ＣＫ）,遮光７５％（ＤＮ）,遮光４０％（ＳＮ）和高、中、低３种水分组合处理,其土壤含水量分别为田间持水量的８５％￣１００％（高水分,ＨＷ）、６５％￣８５％（中水分,ＭＷ）和４５％￣６５％（低水分,ＬＷ）。探讨了土壤水分和遮荫水平共同作用对嘿期至蕾期中午的气孔寻度、净江合速率、蒸腾速率和叶水势等的影响。高水分下ＤＮ和ＳＮ     

Simulating carbon dioxide exchange rates of deciduous tree species: evidence for a general pattern in biochemical changes and water stress response

Background and Aims

Deciduous trees have a seasonal carbon dioxide exchange pattern that is attributed to changes in leaf biochemical properties. However, it is not known if the pattern in leaf biochemical properties – maximum Rubisco carboxylation (V_cmax) and electron transport (J_max) – differ between species. This study explored whether a general pattern of changes in V_cmax, J_max, and a standardized soil moisture response accounted for carbon dioxide exchange of deciduous trees throughout the growing season.

Methods

The model MAESTRA was used to examine V_cmax and J_max of leaves of five deciduous trees, Acer rubrum ‘Summer Red’, Betula nigra, Quercus nuttallii, Quercus phellos and Paulownia elongata, and their response to soil moisture. MAESTRA was parameterized using data from in situ measurements on organs. Linking the changes in biochemical properties of leaves to the whole tree, MAESTRA integrated the general pattern in V_cmax and J_max from gas exchange parameters of leaves with a standardized soil moisture response to describe carbon dioxide exchange throughout the growing season. The model estimates were tested against measurements made on the five species under both irrigated and water-stressed conditions.

Key Results

Measurements and modelling demonstrate that the seasonal pattern of biochemical activity in leaves and soil moisture response can be parameterized with straightforward general relationships. Over the course of the season, differences in carbon exchange between measured and modelled values were within 6–12 % under well-watered conditions and 2–25 % under water stress conditions. Hence, a generalized seasonal pattern in the leaf-level physiological change of V_cmax and J_max, and a standardized response to soil moisture was sufficient to parameterize carbon dioxide exchange for large-scale evaluations.

Conclusions

Simplification in parameterization of the seasonal pattern of leaf biochemical activity and soil moisture response of deciduous forest species is demonstrated. This allows reliable modelling of carbon exchange for deciduous trees, thus circumventing the need for extensive gas exchange experiments on different species.Key words: Carbon budget, deciduous trees, modelling, MAESTRA, soil moisture, species response, transpiration, Acer rubrum, Betula nigra, Quercus nuttallii, Q. phellos, Paulownia elongata     

Allocation of biomass and photoassimilates in juvenile plants of six Patagonian species in response to five water supply regimes

Background and Aims

The growth–differentiation balance hypothesis (GDBH) states that there is a physiological trade-off between growth and secondary metabolism and predicts a parabolic effect of resource availability (such as water or nutrients) on secondary metabolite production. To test this hypothesis, the response of six Patagonian Monte species (Jarava speciosa, Grindelia chiloensis, Prosopis alpataco, Bougainvillea spinosa, Chuquiraga erinacea and Larrea divaricata) were investigated in terms of total biomass and resource allocation patterns in response to a water gradient.

Methods

One-month-old seedlings were subjected to five water supply regimes (expressed as percentage dry soil weight: 13 %, 11 %, 9 %, 7 % or 5 % – field water capacity being 15 %). After 150 d, plants were harvested, oven-dried and partitioned into root, stem and leaf. Allometric analysis was used to correct for size differences in dry matter partitioning. Determinations of total phenolics (TP), condensed tannins (CT), nitrogen (N) and total non-structural carbohydrates (TNC) concentrations were done on each fraction. Based on concentrations and biomass data, contents of TP and CT were estimated for whole plants, and graphical vector analysis was applied to interpret drought effect.

Key Results

Four species (J. speciosa, G. chiloensis, P. alpataco and B. spinosa) showed a decrease in total biomass in the 5 % water supply regime. Differences in dry matter partitioning among treatments were mainly due to size variation. Concentrations of TP, CT, N and TNC varied little and the effect of drought on contents of TP and CT was not adequately predicted by the GDBH, except for G. chiloensis.

Conclusions

Water stress affected growth-related processes (i.e. reduced total biomass) rather than defence-related secondary metabolism or allocation to different organs in juvenile plants. Therefore, the results suggest that application of the GDBH to plants experiencing drought-stress should be done with caution, at least for Patagonian Monte species.     

Plant morphometric traits and climate gradients in northern China: a meta-analysis using quadrat and flora data

Background and Aims

The collection of field data on plant traits is time consuming and this makes it difficult to examine changing patterns of traits along large-scale climate gradients. The present study tests whether trait information derived from regional floras can be used in conjunction with pre-existing quadrat data on species presence to derive meaningful relationships between specific morphometric traits and climate.

Methods

Quadrat records were obtained for 867 species in 404 sites from northern China (38–49°N, 82–132°E) together with information on the presence/absence of key traits from floras. Bioclimate parameters for each site were calculated using the BIOME3 model. Principal component analysis and correlation analysis were conducted to determine the most important climate factors. The Akaike Information Criterion was used to select the best relationship between each trait and climate. Canonical correspondence analysis was used to explore the relationships between climate and trait occurrence.

Key Results

The changing abundance of life form, leaf type, phenology, photosynthetic pathway, leaf size and several other morphometric traits are determined by gradients in plant-available moisture (as measured by the ratio of actual to potential evapotranspiration: α), growing-season temperature (as measured by growing degree-days on a 0 ° base: GDD₀) or a combination of these. Different plant functional types (PFTs, as defined by life form, leaf type and phenology) reach maximum abundance in distinct areas of this climate space: for example, evergreen trees occur in the coldest, wettest environments (GDD₀ < 2500 °Cd, α > 0·38), and deciduous scale-leaved trees occur in drier, warmer environments than deciduous broad-leaved trees. Most leaf-level traits show similar relationships with climate independently of PFT: for example, leaf size in all PFTs increases as the environment becomes wetter and cooler. However, some traits (e.g. petiole length) display different relationships with climate in different PFTs.

Conclusions

Based on presence/absence species data and flora-based trait assignments, the present study demonstrates ecologically plausible trends in the occurrence of key plant traits along climate gradients in northern China. Life form, leaf type, phenology, photosynthetic pathway, leaf size and other key traits reflect climate. The success of these analyses opens the possibility of using quadrat- and flora-based trait analyses to examine climate–trait relationships in other regions of the world.     

Enhancement of artemisinin concentration and yield in response to optimization of nitrogen and potassium supply to Artemisia annua

Background and Aims

The resurgence of malaria, particularly in the developing world, is considerable and exacerbated by the development of single-gene multi-drug resistances to chemicals such as chloroquinone. Drug therapies, as recommended by the World Health Organization, now include the use of antimalarial compounds derived from Artemisia annua – in particular, the use of artemisinin-based ingredients. Despite our limited knowledge of its mode of action or biosynthesis there is a need to secure a supply and enhance yields of artemisinin. The present study aims to determine how plant biomass can be enhanced while maximizing artemisinin concentration by understanding the plant''s nutritional requirements for nitrogen and potassium.

Methods

Experiments were carried out, the first with differing concentrations of nitrogen, at 6, 31, 56, 106, 206 or 306 mg L⁻¹ being applied, while the other differing in potassium concentration (51, 153 or 301 mg L⁻¹). Nutrients were supplied in irrigation water to plants in pots and after a growth period biomass production and leaf artemisinin concentration were measured. These data were used to determine optimal nutrient requirements for artemisinin yield.

Key Results

Nitrogen nutrition enhanced plant nitrogen concentration and biomass production successively up to 106 mg N L⁻¹ for biomass and 206 mg N L⁻¹ for leaf nitrogen; further increases in nitrogen had no influence. Artemisinin concentration in dried leaf material, measured by HPLC mass spectroscopy, was maximal at a nitrogen application of 106 mg L⁻¹, but declined at higher concentrations. Increasing potassium application from 51 to 153 mg L⁻¹ increased total plant biomass, but not at higher applications. Potassium application enhanced leaf potassium concentration, but there was no effect on leaf artemisinin concentration or leaf artemisinin yield.

Conclusions

Artemisinin concentration declined beyond an optimal point with increasing plant nitrogen concentration. Maximization of artemisinin yield (amount per plant) requires optimization of plant biomass via control of nitrogen nutrition.Key words: Artemisia, fertigation, malaria, nitrogen, nutrition, potassium     

Optimal photosynthetic use of light by tropical tree crowns achieved by adjustment of individual leaf angles and nitrogen content

Background and Aims

Theory for optimal allocation of foliar nitrogen (ONA) predicts that both nitrogen concentration and photosynthetic capacity will scale linearly with gradients of insolation within plant canopies. ONA is expected to allow plants to efficiently use both light and nitrogen. However, empirical data generally do not exhibit perfect ONA, and light-use optimization per se is little explored. The aim was to examine to what degree partitioning of nitrogen or light is optimized in the crowns of three tropical canopy tree species.

Methods

Instantaneous photosynthetic photon flux density (PPFD) incident on the adaxial surface of individual leaves was measured along vertical PPFD gradients in tree canopies at a frequency of 0·5 Hz over 9–17 d, and summed to obtain the average daily integral of PPFD for each leaf to characterize its insolation regime. Also measured were leaf N per area (N_area), leaf mass per area (LMA), the cosine of leaf inclination and the parameters of the photosynthetic light response curve [photosynthetic capacity (A_max), dark respiration (R_d), apparent quantum yield (ϕ) and curvature (θ)]. The instantaneous PPFD measurements and light response curves were used to estimate leaf daily photosynthesis (A_daily) for each leaf.

Key Results

Leaf N_area and A_max changed as a hyperbolic asymptotic function of the PPFD regime, not the linear relationship predicted by ONA. Despite this suboptimal nitrogen partitioning among leaves, A_daily did increase linearly with PPFD regime through co-ordinated adjustments in both leaf angle and physiology along canopy gradients in insolation, exhibiting a strong convergence among the three species.

Conclusions

The results suggest that canopy tree leaves in this tropical forest optimize photosynthetic use of PPFD rather than N per se. Tropical tree canopies then can be considered simple ‘big-leaves’ in which all constituent ‘small leaves’ use PPFD with the same photosynthetic efficiency.Key words: Optimal resource allocation, nitrogen, photosynthetic capacity, leaf mass per area, tropical trees, radiation use efficiency, scaling, leaf angle, canopy architecture, big leaf model     

Predator satiation and recruitment in a mast fruiting monocarpic forest herb

Background and Aims

Cross-pollination and satiation of seed predators are often invoked to explain synchronous mast reproduction in long-lived plants. However, explanations for the synchronous death of parent plants are elusive. The roles of synchronous seeding and post-reproductive mortality were investigated in a perennial monocarpic herb (Isoglossa woodii) in coastal dune forest in South Africa.

Methods

Pre-dispersal seed predation and seed production were assessed by measuring fruit and seed set of inflorescences sprayed with insecticide or water and with no spray treatments. Seed predation was measured at different densities of I. woodii plants by monitoring removal rates of seed from the forest floor. The influence of adult plants on establishment of I. woodii seedlings was assessed by monitoring growth and survivorship of seedlings in caged and uncaged 1 × 1 m plots in understorey gaps and thickets.

Key Results

Fruit and seed set were similar between spray treatments. An I. woodii stem produced 767·8 ± 160·8 seeds (mean ± s.e.) on dune crests and 1359·0 ± 234·4 seeds on the foredune. Seed rain was greater on the foredune than in other topographic locations. Seed predation rates were 32 and 54 % on dune crests and in dune slacks, respectively, and decreased with seed abundance, number of inflorescences per stem and plant height. Seedling recruitment was greater beneath synchronously dying adult plants than in natural understorey gaps (no I. woodii). However, seedling growth rate beneath I. woodii mid-way through its life-cycle was less than in gaps, although survivorship was similar.

Conclusions

The selective advantage of masting in I. woodii derives from satiation of both pre- and post-dispersal seed predators. In addition, post-seeding mortality of adult plants facilitates seedling establishment. Satiation of seed predators and the benefits of seedling establishment are strong drivers of the evolution of synchronous monocarpy in I. woodii.