Seasonal variation in standing crop,density and leaf growth rate of the seagrass, Heterozostera tasmanica, in western Port and Port Phillip Bay,Victoria, Australia

Standing crop, density and leaf growth rate of Heterozostera tasmanica (Martens ex Aschers.) den Hartog along with light, temperature, nutrient and sediment characteristics were determined monthly for fifteen months at three study sites in Western Port and one site in Port Phillip Bay, Victoria, Australia. Erect vegetative stems of H. tasmanica were frequently branched, were present throughout the year and accounted for 25–60% of the above-sediment biomass, with the stem proportion higher during winter than summer. At three of the four sites there was a unimodal seasonal pattern in which minimum leaf standing crop (27–61 g dry wt. m^?2), density (600–2000 leaf cluster m^?2) and leaf productivity (0.34–0.77 g dry wt. m^?2 day^?1) generally occurred during winter (June–August) and maximum leaf standing crop (105–173 g dry wt. m^?2), density (2700–5000 leaf cluster m^?2) and leaf productivity (2.6–4.2 g dry wt. m^?2 day^?1) occurred during summer (December–February). A bimodal seasonal pattern with minimum standing crop and density during midsummer occurred at one site. This anomalous seasonal pattern may be due to exposure and desiccation stress during spring low tides. At the site receiving the lowest irradiance, standing crop, density and annual leaf production also were lowest, but length and width of leaves, shoot height and leaf growth rate per leaf cluster were the highest of the four study sites. On average, each leaf cluster at any one of the study sites produced 30–31 leaves per year with mean leaf turnover rates of 1.3–1.7% day^?1. Annual leaf production of H. tasmanica ranged from 410 to 640 g dry wt.m^?2 at the four sites.     

Biomass accumulation and shading effects of epiphytes on leaves of the seagrass, Heterozostera tasmanica, in Victoria,Australia

A method is described for estimating the rate of accumulation of epiphyte biomass on leaves of the seagrass, Heterozostera tasmanica (Martens ex Aschers.) den Hartog and for estimating the effect of epiphyte biomass on photosynthesis of the seagrass. Epiphyte biomass was determined by comparison of the weight per unit area of epiphyte-covered and epiphyte-free leaf blades. Epiphyte weight increased as age of the seagrass leaves increased. Linear regression on epiphyte biomass vs. leaf age estimated the rate of biomass accumulation. Rates varied from 5.7 to 104 μg epiphyte dry weight per cm² of leaf surface per day at three sites in Western Port and Port Phillip Bay, Victoria. Rates of accumulation of epiphyte biomass were generally higher during December through March (summer) than in May (autumn), August (winter) or October (Spring). Light attenuation by epiphytes increase linearly with biomass. The rate of biomass accumulation of epiphytes was compared with leaf growth rate, ambient photon flux density in H. tasmanica beds and the photosynthesis—photon flux density curve of H. tasmanica. This comparison demonstrated that epiphyte biomass can accumulate fast enough to shade H. tasmanica leaves and significantly reduce the time (to less than one half of the leaf life span) in which positive net photosynthesis of the leaf blade is possible.     

Steady-state and dynamic photosynthetic performance and nitrogen partitioning in the shade-demanding plant Panax notoginseng under different levels of growth irradiance

In this study, we examined steady-state and dynamic photosynthetic performance and leaf nitrogen (N) partitioning in the typical shade-demanding herb Panax notoginseng grown along a light gradient. Gas exchange on a leaf area basis was significantly reduced under low irradiance, with gas exchange on a leaf mass basis reaching a maximum value and then decreasing along the light gradient. Specific leaf area significantly increased with decreasing irradiance levels (P < 0.001), whereas carboxylation efficiency was decreased (P < 0.001). In addition, decreasing growth irradiance levels led to declines in maximum carboxylation rate (V _cmax) and maximum electron transport rate (J _max), although V _cmax/mass and J _max/mass were relatively less affected than V _cmax/area and J _max/area. Slow photosynthetic response to simulated sunflecks was observed under low levels of growth irradiance, with stomatal limitations only detected in leaves grown under low-light conditions. Chlorophyll content increased significantly with decreasing irradiance levels. N content on a leaf mass basis apparently increased, while N content on a leaf area basis markedly decreased. The fraction of leaf N allocated to light-harvesting components increased significantly with decreasing growth irradiance levels, whereas the fraction allocated to carboxylation and bioenergetics was significantly reduced. As an adaptation strategy to growth irradiance, we conclude that adjustments in specific leaf area may be more important than changes in leaf physiology and biochemistry in typical shade-demanding species such as P. notoginseng.     

Heterozostera tasmanica (Martens ex aschers.) den Hartog in Chile

A 1.2 km² meadow of Heterozostera tasmanica (Martens ex Aschers.) den Hartog was found at Pto. Aldea, Bahia Tongoy, 60 km south of Coquimbo, 30°S on the coast of Chile. This provides documented evidence of seagrass occurrence on the west coast of South America. Since H. tasmanica is widespread throughout southern Australia, the Chilean stock represents a highly disjunct population. It appears from several growth parameters (density, leaf biomass, leaf area index, specific leaf area, δ ¹³C) that the H. tasmanica meadow at Pto. Aldea is very similar to those found in other parts of the world.     

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.     

Biomass allocation,morphology and photosynthesis of invasive and noninvasive exotic species grown at four irradiance levels

We tested the hypotheses that invasive species had higher irradiance plasticity, capture ability and efficiency than noninvasive species using two invasive aliens – Ageratina adenophora and Chromolaena odorata, and one noninvasive alien – Gynura sp. The three aliens were grown at 4.5%, 12.5%, 36%, 50% and 100% irradiances for 64 days before harvesting. The plastic response of specific leaf area (SLA) contributed to improved light interception at low irradiance, carbon gain and water balance at high irradiance. It was a good predictor for intraspecific irradiance responses of leaf area ratio (LAR), leaf area:root mass ratio, maximum photosynthetic rate (P_max) and net assimilation rate (NAR). Biomass allocation-related traits were species specific and their plasticity to irradiance was low. The high root mass fraction, leaf mass fraction and LAR distinguished the two invaders from Gynura. However, other resource capture-related traits, such as SLA, NAR and P_max, were not always higher for the invaders than for Gynura. Furthermore, plasticity to irradiance was not different between the invasive and noninvasive aliens. With increasing irradiance, Gynura decreased biomass investment to roots and leaves but increased the investment to support structures adversely affecting both low and high irradiance acclimation. Ageratina might invade new habitat successfully through tolerating shading at low irradiance and outshading competitors by forming dense stands when irradiance is increased. The results suggested that both resource capture-related traits and irradiance acclimation conferred competitive advantage to the two invaders and some traits were common for invasive and noninvasive aliens but others were specific for invaders.     

Growth rates and assimilate partitioning in the elongation zone of tall fescue leaf blades at high and low irradiance

Tall fescue (Festuca arundinacea Schreb.) leaf blades elongated 33% faster at continuous low than at continuous high irradiance (60 versus 300 micromoles per second per square meter photosynthetic photon flux density) when temperature of the leaf elongation zone was held constant at 21°C. Increased rate of elongation was associated with a near proportional increase in length of the elongation zone (+38%). In contrast, growth in width and thickness was decreased at low irradiance, resulting in only a 12% increase in leaf area production and 5% less total growth-associated water deposition than at high irradiance. At low irradiance dry matter (DM) import into the elongation zone was 28% less, and 55% less DM was used per unit leaf area produced. DM use in synthesis of structural components (i.e. DM less water-soluble carbohydrates) was only 13% less at low irradiance, whereas water-soluble carbohydrates (WSC) deposition was 43% less. The lower rate of WSC deposition at low irradiance was associated with a higher net rate of monosaccharide deposition (+39%), whereas net deposition rates for sucrose (−27%) and fructan (−56%) were less than at high irradiance. Still, at low irradiance, net fructan accumulation accounted for 64% of WSC deposition, i.e. 25% of DM import, demonstrating the high sink strength of the leaf elongation zone.     

Spatio-temporal variability in the photosynthetic characteristics of Zostera tasmanica measured by PAM

Rapid light curves (RLCs), based on pulse amplitude modulated (PAM) fluorometry, were used to investigate the spatio-temporal variability in photosynthesis versus irradiance parameters (α, I_k and P_max) and the F_v/F_m ratio of the seagrass Zostera tasmanica (formerly Heterozostera tasmanica). Spatial variation was examined across scales ranging from within a leaf (cms) to across the bed (ms), using a nested analysis of covariance sampling design. Overall, significant variation was identified at all scales examined, excluding the largest scale (area). Patterns of variability differed among individual parameters; however a high percentage of the variation was consistently assigned to the covariates, age (within and between leaves) for all parameters, except P_max.     

Photosynthetic responses of <Emphasis Type="Italic">Chrysanthemum morifolium</Emphasis> to growth irradiance: morphology,anatomy and chloroplast ultrastructure

Seedlings of Chrysanthemum, cultivar ‘Puma Sunny’, were grown under a range of shading regimes (natural full sunlight, 55, 25, and 15% of full sunlight) for 18 days. Here, we characterized effects of varying light regimes on plant morphology, photosynthesis, chlorophyll fluorescence, anatomical traits, and chloroplast ultrastructure. We showed that leaf color was yellowish-green under full sunlight. Leaf area, internode length, and petiole length of plants were the largest under 15% irradiance. Net photosynthetic rate, water-use efficiency, PSII quantum efficiency, and starch grain were reduced with decreasing irradiance from 100 to 15%. Heavy shading resulted in the partial closure of PSII reaction centers and the CO₂ assimilation was restricted. The results showed the leaves of plants were thinner under 25 and 15% irradiance with loose palisade tissue and irregularly arranged spongy mesophyll cells, while the plants grown under full sunlight showed the most compact leaf palisade parenchyma. Irradiance lesser than 25% of full sunlight reduced carbon assimilation and led to limited plant growth. Approximately 55% irradiance was suggested to be the optimal for Chrysanthemum morifolium.     

荆条叶性状对野外不同光环境的表型可塑性

光照是影响植物生长和分布的重要环境因子。对生长在野外5种不同光环境下(林外、阔叶林林缘、阔叶林林下、针叶林林窗和针叶林林下)的荆条的叶片进行取样研究,通过对光合作用光响应曲线、叶绿素荧光、叶绿素含量、叶片氮磷含量以及叶片形态的测量,来反映荆条对不同光环境的表型可塑性。研究结果表明,荆条叶片对于野外不同的光环境具有很好的适应机制,叶片功能性状受到结构性状的调节。低光下通过高的比叶面积(SLA)、单位质量叶绿素含量、光系统II最大量子产量,低的暗呼吸速率、光饱和点、光补偿点、叶绿素a,b的比值来提高对光能的利用效率,维持生长;高光下则通过与SLA有关的叶片结构的变化对光合作用进行调节。大多数的叶性状只受到日光照总量的影响,SLA的大小与日最高光强有关,可以对不同日变化模式的光照做出迅速的响应,是适应不同光照的敏感指标。尽管光照是不同光环境下影响荆条叶性状的主要环境因子,土壤养分含量同样会对叶性状产生影响,高土壤养分下的高叶长与叶柄长的比值体现了植物对资源获取和支撑结构之间分配的权衡。     

The reproductive cycle of the asteroid Coscinasterias muricata in Port Phillip Bay, Victoria, Australia

The reproductive cycles of two populations of Coscinasterias muricata from Port Phillip Bay, Victoria, Australia are described in terms of organ indices, oocyte development and progesterone levels. Both Governor's Reef and South Channel Fort populations exhibited clearly defined reproductive cycles with two spawning periods, during summer and during spring. In both populations, the pyloric caecal index and gonadal index showed inverse cycles suggesting nutrient translocation from the pyloric caeca to the gonads for gametogenesis. Physiological changes, such as decreases in pyloric caecal index, and increases in gonadal index and oocyte diameter were observed following the months of increased progesterone concentrations in the female pyloric caeca. These results indicate a role for this steroid in the reproductive cycle. The results from the Governor's Reef population suggest that the influence of progesterone on the reproductive cycle of female C. muricata is regulated by photoperiod. Results also provide evidence of a role for seawater temperature in the reproductive process of this species.     

Effects of irradiance on growth and flowering in the shade plant, cineraria

Effects of total irradiance on growth and flowering were studied in cineraria cv. Cindy Blue grown under warm (mean 21°C) glasshouse conditions. Efficiency of light conversion for leaf and shoot dry weight increase were reduced from 0.08 to 0.02 as the mean daily light integral increased from 0.9 to 4.4 MJ m^-2 day^-1 but no significant difference in leaf area were associated with this. Specific leaf area decreased exponentially from 0.07 to 0.02 m²g¹ over the cumulative irradiance range 23 to 127 MJ m^-2 after the start of treatments and thereafter remained stable. A light integral of 19.2 MJ m^-2 were required for initiation of one leaf in plants grown under a daily integral of 4.4 MJ m^-2 day^-1, as compared with only 5.1 MJ m^-2day^-1 required per leaf in plants grown at less than 0.9 MJ m^-2day^-1. Neither chronological duration of juvenile development nor leaf number below the flower was affected by irradiance. However, as the rate of leaf initiation increased with irradiance up to 2.4 MJ m^-2day^-1 so the rate of progress to flower visibility increased linearly with irradiance over the same range. This rate then remained constant from 2.4 to 4.4 MJ m^-2day^-1. Length of the main flowering shoot decreased and the number of flowering shoots increased as irradiance increased from 0.9 to 2.4 MJ m^-2 day^-1 and then remained unchanged by further increases in irradiance.     

Leaf orientation,photorespiration and xanthophyll cycle protect young soybean leaves against high irradiance in field

In order to fully understand the adaptive strategies of young leaves in performing photosynthesis under high irradiance, leaf orientation, chloroplast pigments, gas exchange, as well as chlorophyll a fluorescence kinetics were explored in soybean plants. The chlorophyll content and photosynthesis in young leaves were much lower than that in fully expanded leaves. Both young and fully expanded leaves exhibited down-regulation of the maximum quantum yield (F_V/F_M) at noon in their natural position, no more serious down-regulation being observed in young leaves. However, when restraining leaf movement and vertically exposing the leaves to 1200 μmol m⁻² s⁻¹ irradiance, more pronounced down-regulation of F_V/F_M was observed in young leaves; and the actual photosystem II (PS II) efficiency (Ф_PSII) drastically decreased with the significant enhancement of non-photochemical quenching (NPQ) and ‘High energy’ quenching (q_E) in young leaves. Under irradiance of 1200 μmol m⁻² s⁻¹, photorespiration (P_r) in young leaves measured by gas exchange were obviously lower, whereas the ratio of photorespiration/gross photosynthetic rate (P_r/P_g) were higher than that in fully expanded leaves. Compared with fully expanded leaves, young leaves exhibited higher xanthophyll pool and a much higher level of de-epoxidation components when exposure to high irradiance. During leaf development, the petiole angle gradually increased all the way. Especially, the midrib angle decreased with the increasing of irradiance in young leaves; however, no distinct changes were observed in mature leaves. The changes of leaf orientation greatly reduced the irradiance on young leaf surface under natural positions. In this study, we suggested that the co-operation of leaf angle, photorespiration and thermal dissipation depending on xanthophyll cycle could successfully prevent young leaves against high irradiance in field.     

Effects of light regime on the growth,leaf morphology,and water relations of seedlings of two species of tropical trees

An experiment was conducted with Heliocarpus appendiculatus, a pioneer or large gap species of tropical moist forest in Costa Rica, and Dipteryx panamensis, a small gap species. Seedlings were grown in full sun, partial (80%) shade, and full (98%) shade. After one month of growth they were switched between environments and grown for two more months. Growth in height of Heliocarpus was greatly affected by irradiance, being increased in response to full shade and decreased in full sun. Height of Dipteryx was unaffected by irradiance level. Survival of Heliocarpus seedlings was only 49% in full shade, whereas Dipteryx had 100% survival. Biomass of Heliocarpus was not significantly greater in full sun than in partial shade whereas it was for Dipteryx. The response of root: shoot ratio was similar for both species. They were lowest in full shade and highest in full sun. Heliocarpus exhibited greater changes in leaf thickness, specific leaf weight, and stomatal density than did Dipteryx. Stomatal conductance of both species was lower in full shade and full sun than in partial shade. The results of the experiment indicate that growth of Heliocarpus is more plastic than that of Dipteryx in response to changes in irradiance. Previous environment did not affect the response to the present environment in either species. Both species responded positively to increases in irradiance.     

Responses of root length/leaf area ratio and specific root length of an understory herb, Pteridophyllum racemosum, to increases in irradiance

The understory evergreen perennial Pteridophyllum racemosum Sieb. et Zucc. (Papaveraceae) has the ability to increase root mass per unit transpiring leaf area (RMA) if irradiance increases gradually over several years. In this study, we examined how P. racemosum changes its root length/leaf area ratio and specific root length when the species encounters abrupt increases in irradiance, such as sudden and unexpected canopy openings. Plants were transplanted from a low light condition in a subalpine wave-regenerating forest (photon flux density on the forest floor relative to the full sun (RPFD) was 2.7%) to a high light condition in a glasshouse (30% RPFD) (LH treatment). Transplantation from the low light condition in the forest to a low light condition in the glasshouse (LL) and transplantation from a high light condition in the forest (33% RPFD) to a high light condition in the glasshouse (HH) were also conducted as controls. Compared to the LL plants, the LH plants exhibited significant increases in RMA and root length/leaf area ratio from 30 to 70 days after transplantation. On the other hand, the effect of increased irradiance on specific root length (SRL) was weak, and both the LL and LH plants showed increased SRL from 30 to 70 days after transplantation. Increased SRL results from longer root length per unit construction cost. We concluded that increased root length/leaf area ratio of P. racemosum in response to abrupt increases in irradiance was caused by a combination of enhanced carbon allocation to roots with increased SRL.     

Interaction effects of phosphorus and zinc on photosynthesis, growth and yield of dwarf bean grown in two environments

Two experiments were conducted in a factorial combination of three Zn levels (0, 10 and 40 mg Zn kg^-1 soil) and two P levels (0 and 200 mg P kg^-1 soil). Experiment 1 was carried out during winter in a heated glasshouse, and experiment 2 during summer under a rain shelter. Plants of dwarf bean (Phaseolus vulgaris L., cv. Borlotto nano) were grown in pots filled with sandy soil. In both experiments, leaf Zn concentration was reduced by the addition of P to plants grown at low Zn supply. However, leaf Zn concentration lower than the critical level was observed only during experiment 2, and the main effects of low Zn were reductions of internode length, light use efficiency and maximum photosynthetic rate. In plants with leaf Zn concentration lower than the critical level, saturating irradiance levels fell from _∼1000 μmol m^-2 s^-1 PPFD to _∼300–400 μmol m^-2 s^-1 PPFD. Reduction of net photosynthesis was observed from the beginning of flowering and led to decreased seed production. This revised version was published online in June 2006 with corrections to the Cover Date.     

Regulation of tomato fruit ascorbate content is more highly dependent on fruit irradiance than leaf irradiance

Background and Aims

The mechanisms involving light control of vitamin C content in fruits are not yet fully understood. The present study aimed to evaluate the impact of fruit and leaf shading on ascorbate (AsA) accumulation in tomato fruit and to determine how fruit sugar content (as an AsA precursor) affected AsA content.

Methods

Cherry tomato plants were grown in a glasshouse. The control treatment (normally irradiated fruits and irradiated leaves) was compared with the whole-plant shading treatment and with leaf or fruit shading treatments in fruits harvested at breaker stage. In a second experiment, the correlation between sugars and AsA was studied during ripening.

Key Results

Fruit shading was the most effective treatment in reducing fruit AsA content. Under normal conditions, AsA and sugar content were correlated and increased with the ripening stage. Reducing fruit irradiance strongly decreased the reduced AsA content (−74 %), without affecting sugars, so that sugar and reduced AsA were no longer correlated. Leaf shading delayed fruit ripening: it increased the accumulation of oxidized AsA in green fruits (+98 %), whereas it decreased the reduced AsA content in orange fruits (−19 %), suggesting that fruit AsA metabolism also depends on leaf irradiance.

Conclusions

Under fruit shading only, the absence of a correlation between sugars and reduced AsA content indicated that fruit AsA content was not limited by leaf photosynthesis or sugar substrate, but strongly depended on fruit irradiance. Leaf shading most probably affected fruit AsA content by delaying fruit ripening, and suggested a complex regulation of AsA metabolism which depends on both fruit and leaf irradiance and fruit ripening stage.Key words: Ascorbate, fruit quality, irradiance, shading, Solanum lycopersicon, sugars, tomato, vitamin C     

Effect of irradiance during growth of Glycine max on photosynthetic capacity and percent activation of ribulose 1,5-bisphosphate carboxylase

The initial (in vivo) and total (activity present after preincubation with CO₂ and Mg²⁺) activities of ribulose bisphosphate carboxylase were both assayed in extracts of leaves of soybean (Glycine max) plants which had been grown under 4 different irradiance levels. The total carboxylase activity per unit leaf area decreased with decreased irradiance during growth but was not different on a dry weight basis. The initial activity as a percentage of the total activity was unchanged (approximately 95%) except in leaves of plants grown at the lowest irradiance (74%). When the plants grown at the lowest irradiance were exposed to high irradiance, the initial activity was increased to 93% of the total. Light saturated rates of photosynthesis per unit leaf area were lower and saturated at lower irradiance for plants grown at lower irradiances. Initial carboxylase activity was correlated closely (r²=0.84) with leaf photosynthesis rate on a dry weight basis.     

Effects of light intensity on leaf photosynthetic characteristics,chloroplast structure,and alkaloid content of <Emphasis Type="Italic">Mahonia bodinieri</Emphasis> (Gagnep.) Laferr.

The variation of light intensity has obvious effects on leaf external morphology, internal anatomy, and physiological characteristics; it even induces changes in secondary metabolite production. The effects of different irradiance levels on biomass, gas exchange parameters, and photosynthetic pigment contents in Mahonia bodinieri (Gagnep.) Laferr. were analyzed here. Combined analyses of physiology, cytology, and HPLC were used to study the differences in leaf morphology, structure, physiological characters, and alkaloid content in response to different irradiances. The results indicated that the highest foliar biomass was observed under I ₅₀ (50 % of full sunlight) followed by I ₃₀ (30 % of full sunlight), the highest net photosynthetic rate, stomatal conductance, transpiration rate values were observed under I ₃₀ followed by I ₅₀, and lower values occurred in I ₁₀ (10 % of full sunlight) and I ₁₀₀ (full sunlight). With increased light intensity, total leaf area and the contents of chlorophyll a (Chl a), chlorophyll b (Chl b), and chlorophyll (Chl a+b) per unit leaf area were clearly reduced, whereas leaf mass per area, carotenoid content, leaf thickness, thickness of palisade and spongy parenchyma, and stomatal density were all significantly increased. Electron microscopic observation revealed that the number of grana, stroma lamellae and the number of starch grains in chloroplasts were decreased, the number of plastoglobuli was increased when irradiance levels increased. The estimated total yield of alkaloids in a single plant was higher under I ₃₀ and I ₅₀ than under I ₁₀ or I ₁₀₀ as a result of the higher biomass of the plants. Therefore, I ₃₀ and I ₅₀ were not only beneficial to increase biomass, but also suitable for the synthesis and accumulation of the major secondary metabolites (alkaloids). Our findings provide valuable data for the determination and regulation of irradiance levels during artificial cultivation of M. bodinieri.