Pasture intake and milk production of dairy cows rotationally grazing on multi-species swards

Increasing plant species diversity has been proposed as a means for enhancing annual pasture productivity and decreasing seasonal variability of pasture production facing more frequent drought scenarios due to climate change. Few studies have examined how botanical complexity of sown swards affects cow performance. A 2-year experiment was conducted to determine how sward botanical complexity, from a monoculture of ryegrass to multi-species swards (MSS) (grasses-legumes-forb), affect pasture chemical composition and nutritive value, pasture dry matter (DM) intake, milk production and milk solids production of grazing dairy cows. Five sward species: perennial ryegrass (L as Lolium), white clover and red clover (both referred to as T as Trifolium because they were always sown together), chicory (C as Cichorium) and tall fescue (F as Festuca) were assigned to four grazing treatments by combining one (L), three (LT), four (LTC) or five (LTCF) species. Hereafter, the LT swards are called mixed swards as a single combination of ryegrass and clovers, whereas LTC and LTCF swards are called MSS as a combination of at least four species from three botanical families. The experimental area (8.7 ha) was divided into four block replicates with a mineral nitrogen fertilisation of 75 kg N/ha per year for each treatment. In total, 13 grazing rotations were carried out by applying the same grazing calendar and the same pasture allowance of 19 kg DM/cow per day above 4 cm for all treatments. Clover represented 20% of DM for mixed and MSS swards; chicory represented 30% of DM for MSS and tall fescue represented 10% of DM for LTCF swards. Higher milk production (+1.1 kg/day) and milk solids production (+0.08 kg/day) were observed for mixed swards than for ryegrass swards. Pasture nutritive value and pasture DM intake were unaffected by the inclusion of clover. Pasture DM, organic matter and NDF concentrations were lower for MSS than for mixed swards. Higher milk production (+0.8 kg/day), milk solids production (+0.04 kg/day) and pasture DM intake (+1.5 kg DM/day) were observed for MSS than for mixed swards. These positive effects of MSS were observed for all seasons, but particularly during summer where chicory proportion was the highest. In conclusion, advantages of grazing MSS on cow performance were due to the cumulative effect of improved pasture nutritive value and increased pasture DM intake that raised milk production and milk solids production.     

Acclimation of Myrtus communis to contrasting Mediterranean light environments - effects on structure and chemical composition of foliage and plant water relations

Leaf anatomical and chemical characteristics, water relations and stomatal regulation were studied in the shrub Myrtus communis growing under two contrasting Mediterranean light environments (full light versus 30% of full light) during the spring-summer period. These studies aimed to assess plant response to the combined effects of light and water availability. Foliar morphology, anatomy and chemistry composition acclimated positively to light conditions. Leaves of sun-exposed plants were thicker (38.7%) than those of shaded plants, mainly due to increased palisade parenchyma thickness, had a higher nitrogen concentration and stomatal density than the shade ones, which maximized foliar area (>SLA) and Chl/N molar ratio to improve light interception. Chlorophyll concentration per leaf area (Chl(a)) was always higher in sun leaves while, as expressed on dry mass (Chl(m)), significant differences were only apparent in September, shade leaves presenting higher values. During the summer period Chl(a) and Chl(m) markedly declined in sun leaves and remained unchanged in shade ones. The ratio of chlorophyll a/b was not affected either by the light intensity or by the season. Shade leaves presented generally a higher concentration of soluble carbohydrates per dry mass. No significant differences in starch concentration were apparent between sun and shade leaves and a gradual depletion occurred during the water stress period. Maximum stomatal conductances correlated positively with predawn water potential. Throughout the season, sun plants always presented higher leaf conductance to water vapour and lower minimum leaf water potentials, indicating an interaction of light-environment on these water relation parameters. Stomatal closure constitutes a mechanism to cope with diurnal and seasonal water deficits, sun plants presenting a more efficient control of water losses during water deficiency period. In addition, both sun and shade plants evidenced leaf osmotic adjustment ability in response to water stress, which was greater in sun ones.     

Competition between grass and clover in spring as affected by nitrogen fertiliser

Detailed measurements of irrigated ryegrass/white clover swards growing without interruption, with or without nitrogen fertiliser in spring, showed that the relative growth rate of clover was as great as that of grass, in the + N sward, and considerably greater than grass, in the – N sward. Clover leaves were not overtopped by grass leaves. Indeed, in both swards, clover had a greater proportion of its leaf lamina area in the upper, well lit, layers of the canopy than grass did. Consequently, clover had a greater mean rate of leaf photosynthesis in situ in the sward than grass. Clover's advantage in photosynthetic rate per leaf area was offset to some extent by its smaller ratio of leaf area to total above-ground dry weight than grass. The consequences of these results for our understanding of competition between grass and clover in mixed swards are discussed.     

Influence of contrasting availabilities of water and nutrients on the radiation use efficiency in C3 and C4 grasses

The radiation use efficiency (RUE) model is one of the most used tools to generate large spatial and temporal scale net primary productivity (NPP) estimations by remote sensing. It involves two key issues to make accurate estimations of NPP: the estimation of the fraction of photosynthetically active radiation (PAR) intercepted by vegetation (fPAR) and the estimation of the plant RUE. The objectives of this work were to quantify the above‐ground RUE under optimal water and nutrient conditions in two C₃ and one C₄ grass species and to analyse the effect of restrictions in these factors upon RUE by comparing both metabolic pathways. Grasses were cultivated from seeds and four treatments combining contrasting availabilities of water and nutrients were applied. RUE values were calculated from measurements of the incoming PAR, fPAR and productivity. In each of the species, plants with sufficient water and nutrients showed the highest RUE (2.61–3.52 g MJ^?1), whereas those with deficiencies in both resources presented the lowest RUE (1.15–2.39 g MJ^?1). Cynodon dactylon (C₄) was the species with higher value of RUE and no significant differences were detected between treatments. However, no significant differences were detected between C. dactylon and D. glomerata under no stress treatment (N1W1) and between C. dactylon and L. perenne under water stress treatment (N1W0). RUE values of Dactylis glomerata (C₃) diminished if only one of the two stress factors was presented, while Lolium perenne (C₃) only when both stress factors were present. The decreases under stress treatments were between 35% and 60% compared with the no stress treatment. When regional NPP is estimated it is therefore important to take into account the decrease in the RUE, especially in areas under severe stress.     

Effects of solar radiation on bacterial and fungal density on aquatic plant detritus

1. The effects of solar radiation on bacterial and fungal growth on aquatic macrophyte detritus were studied in a microcosm experiment. Senescent leaves of Phragmites australis were incubated for 63 days in shallow water in the shade under photosynthetically active radiation (PAR) together with ultraviolet radiation, or under filters removing either ultraviolet B (UVB) or both UVB and ultraviolet A (UVA). 2. Bacterial abundance and bacterial ³H-leucine incorporation in the water were measured, together with α- and β-D-glucosidase activity. In addition, bacterial abundance and fungal biomass associated with the litter were measured. 3. The results indicate that both PAR and UVA affect the micro-organisms involved in the decomposition of leaf litter. The α/β-D-glucosidase activity ratio was less than one in irradiated and more than one in shaded microcosms, suggesting a change in the substrate dissolved organic matter composition towards more β- than α-glycosidic linkages as a result of solar radiation. 4. Microcosms receiving UVB displayed a significantly higher β-D-glucosidase activity than shaded microcosms, and those exposed to PAR or PAR + UVA, demonstrating the potential importance of UVB radiation. 5. The free-living bacteria tended to be dominated by filamentous forms in microcosms subject to solar radiation, especially PAR, and attached microbial communities showed a greater tendency to be dominated by bacteria in irradiated microcosms than in shaded microcosms.     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

Diurnal variation of photosynthesis and photoinhibition in tea: effects of irradiance and nitrogen supply during growth in the field.

Diurnal changes in the rate of photosynthesis (A) of mature tea (Camellia sinensis (L.) O. Kuntze) bushes grown at high elevation in the field in Sri Lanka, were related to environmental conditions. Bushes were either unshaded, receiving 100% of incident photosynthetically active radiation (PAR), moderately shaded, (65% PAR) or heavily shaded (30% PAR). These treatments were combined with nitrogen fertilizer applications of 0, 360 and 720 kg ha(-1) year(-1). When recently fully expanded leaves were measured under the growing conditions on bright, clear days from dawn to dusk, A was greatest in the morning with increasing radiation between approximately 8 h and 10 h. Stomatal conductances (g(s)) and substomatal carbon dioxide concentrations (C(i)) were then large, leaf temperatures (T(L)) cool, and saturated water vapour deficits (VPD) small. However, as the irradiance, T(L) and VPD increased towards midday, A, g(s), photochemical quenching, and C(i) decreased, and non-photochemical quenching increased. In the late afternoon, irradiance, T(L) and VPD fell, but despite the relatively large increase in g(s) and C(i), A remained low; however, it recovered overnight. The zero-N treatment decreased total-N content of leaves by 50% and A by c. 20% (not significant). Leaves of unshaded plants receiving least N had significantly (P<0.05) smaller A and greater total sugar content than shaded but with abundant N, A and sugars did not differ between shade treatments. Analysis of the responses of A to environment in the morning compared to the afternoon, and of chlorophyll fluorescence, suggests that A was photoinhibited as a consequence of greatly increased PAR, whilst decreasing g(s) (related to changes in PAR, VPD and T(L)) caused C(i) to fall. End-product inhibition of A is not consistent with decreased C(i). Inhibition of A as a result of photoinhibition was minimized, but not eliminated, by abundant N. Interactions between factors regulating A in tea are discussed.     

Irradiance heterogeneity within crown affects photosynthetic capacity and nitrogen distribution of leaves in Cedrela sinensis

Because light conditions in the forest understory are highly heterogeneous, photosynthetic acclimation to spatially variable irradiance within a crown is important for crown‐level carbon assimilation. The effect of variation in irradiance within the crown on leaf nitrogen content and photosynthetic rate was examined for pinnate compound leaves in saplings of Cedrela sinensis, a pioneer deciduous tree. Five shading treatments, in which 0, 25, 50, 75 and 100% of leaves were shaded, were established by artificial heavy shading using shade screen umbrellas with 25% transmittance. Although the nitrogen content of leaves was constant regardless of shading treatment, ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) content and light‐saturated photosynthetic capacity were lower in shade leaves within partially shaded crowns than within fully shaded crowns. Shade leaves within partially shaded crowns contained higher amount of amino acids. Most shade leaves died in partially shaded crowns, whereas more than half of shade leaves survived in totally shaded crowns. Assumptions on photosynthetic acclimation to local light conditions cannot explain why shade leaves have different photosynthetic capacities and survival rates in between partially and totally shaded crowns. Irradiance heterogeneity within the crown causes a distinct variation in photosynthetic activity between sun and shaded leaves within the crown.     

Effect of Previous Defoliation Regime and Mineral Nitrogen on Regrowth in White Clover Swards: Photosynthesis, Respiration, Nitrogenase Activity and Growth

Small swards of white clover (Trifolium repens L.) cv. Haifawere grown in solution culture in a controlled environment at24 °C day/18 °C night and receiving 500 µE m^-2S^–1 PAR during a 14-h photoperiod. The swards were cuteither frequently (10-d regrowth periods) or infrequently (40-dregrowth) over 40 d before being cut to 2 cm in height. Halfof the swards received high levels of nitrate (2–6 mMN in solution every 2 d) after defoliation while the othersreceived none. Changes in d. wt, leaf area and growing pointnumbers were recorded over the following 10 d. CO₂ exchangewas measured independently on shoots and roots and nitrogenase-linkedrespiration was estimated by measuring nodulated root respirationat 21% and 3% oxygen in the root atmosphere. There was a general pattern in all treatments consisting ofan initial d. wt loss from roots and stubble and reallocationto new leaves, followed by a period of total d. wt gain andrecovery, to a greater or lesser extent, of weight in non-photosyntheticparts. Frequently cut swards had a smaller proportion of theirshoot d. wt. removed by cutting and had a greater shoot d. wt,growing point number and leaf area at the start of the regrowthperiod. As a result of these differences, and also because ofdifferences in relative growth rates, frequently cut swardsmade more regrowth than infrequently cut. Initial photosyntheticrates were higher in frequently cut swards, although the laminaarea index was very low, and it was concluded that stolons andcut petioles made a significant contribution to carbon uptakeduring the first few d. Infrequently cut swards continued toallocate carbon to new and thinner leaves at the expense ofroots and stubble for longer than frequently cut swards andas a result achieved a similar lamina area index after 10 d. Nitrogenase-linked respiration was low in all treatments immediatelyafter cutting: frequently cut swards receiving no nitrate maintainedhigh nitrogenase activity, whereas recovery took at least 5d in infrequently cut swards. Swards which received nitrateafter cutting maintained only low rates of nitrogenase-linkedrespiration and their total nodulated root respiration overthe period was lower than those receiving no nitrogen: greaterregrowth in nitrate fed swards over the 10 d compared to N₂-fixingswards was in proportion to this lower respiratory burden. White clover (Trifolium repens L.), defoliation, regrowth, nitrogen, photosynthesis, respiration, nitrogenase-activity     

Chloroplastic Carbon Dioxide Concentration in Norway Spruce (Picea Abies [L.] Karst.) needles relates to the position within the crown

Differences between sun (E) and shaded (S) foliage were studied in a Norway spruce (Picea abies [L.] Karst.) stand. Response curves describing the dependence of the CO2 assimilation rate (PN) on the CO2 concentration at the catalytic site of ribulose-1,5-bisphosphate carboxylase/oxygenase, RuBPCO (PN-Cc) were estimated using the simultaneous measurements of chlorophyll fluorescence and leaf gas exchange. Higher PN, higher electron transport (Ja), higher carboxylation capacity (Vc), and higher RuBPCO activity () for sun acclimated needles was found. The S-needles had higher portion of internal limitation and higher CO2 compensation concentration () than the E-needles. Because higher degree of limitation of photosynthesis by carboxylation was ascertained, it can be assumed that photosynthesis in shade foliage is limited mainly by lower carboxylation capacity and by low chloroplastic CO2 concentration     

Effects of Nitrogen fertilisation and regrowth period on fatty acid concentrations in perennial ryegrass (Lolium perenne L.)

Information on lipids in forages is scarce, and effects of nitrogen (N) application level and regrowth period on the fatty acid (FA) concentration and composition of perennial ryegrass (Lolium perenne L.) were studied. N was applied at 0, 45 and 100 kg ha^?1, and swards were cut after various regrowth periods, resulting in six treatments designed as randomised blocks with three replicates. The stages of development ranged from vegetative to elongating swards, with herbage yield levels from 1.9 to 4.2 t dry matter (DM) ha^?1. Concentrations of individual FA were determined by gas chromatography, and canopy characteristics and herbage quality were assessed. The treatments resulted in canopies with contrasting DM yields and protein concentrations. Five FAs, representing 98% of total FAs, were studied in detail. On an average, the concentration of these major FAs in fresh grass was 15.1 g kg^?1 DM, and 69% of the major FAs consisted of C18:3. Regrowth period affected (P < 0.05) the total FA concentration, and significantly (P < 0.01) lower concentrations of C18:3 and C16:1 were found after a longer period of regrowth. N application resulted in higher (P < 0.001) concentrations of all FAs. The FA composition was not affected by N application, but a longer regrowth period significantly (P < 0.001) decreased the proportion of C18:3 and increased those of C18:2 and C16:0. A strong, positive overall linear relation was found between the concentrations of total FAs and C18:3 with the crude protein concentration in the herbage. These studies demonstrate opportunities to affect the FA concentration and composition of FA in herbage through management strategies, which could affect milk FA composition.     

Population ecology in a natural<Emphasis Type="Italic">syneilesis palmata</Emphasis> stand: II. vegetative growth and population structure

Vegetative growth and changes within populations ofSyneilesis palmata (THUNB>) MAX. Usannamul were monitored in two natural stands with different levels of incoming light. This perennial, shade-tolerant herb was measured for its plant size, petiole length, leaf breadth, and leaflet number over 11 years in a moderately shaded (MS) stand and over 7 years in a severely shaded (SS) stand. At the end of each growing season, a shoot primordium developed at the center of the root system. Correlation coefficients were mostly high between pairings of two properties - total weight, petiole length, petiole weight, leaf breadth, leaf blade weight, leaf area, specific leaf area, and leaflet number. Mean annual mortality of this species was higher in SS (22.97%) than in MS (8.85%), but great fluctuations were seen from year by year. Mortality was lowest for medium-sized plants. Regarding petiole length, differences in mean annual growth rates were conspicuous, i.e., 2.6% in MS and 8.4% in SS, while growth rates for leaf breadth were 3.1% in MS and 24.2% in SS. Changes in plant frequency within individual size classes roughly showed a normal distribution curve. However, the mode varied year by year, and such changes were more remarkable in SS than in MS. Under severe shading,S. palmata had a higher growth rate and but also greater mortality than did plants under moderate shade. Therefore, one can conclude that the population structure in SS was unstable over time.     

From protégé to nurse plant: establishment of thorny shrubs in grazed temperate woodlands

Question: Thorny shrubs play keystone roles in grazed ecosystems by defending non‐protected plants against herbivores, but their establishment in grazed ecosystems is poorly understood. Which factors control establishment of recruits of thorny nurse shrubs in grazed temperate woodlands? Location: Ancient grazed temperate woodlands (52°32′N, 6°36′E), The Netherlands. Methods: We surveyed biotic and abiotic factors for saplings of thorny nurse shrubs in plots with and without saplings. To disentangle these factors, we performed a transplantation experiment over two growing seasons with nurse shrub saplings (Prunus spinosa and Crataegus monogyna) planted in two dominant vegetation types – tall unpalatable swards and short grazed lawns – half of them protected from herbivory via exclosures. Results: Plots with shrub saplings had taller surrounding vegetation, higher soil pH and higher soil moisture than plots without saplings. These plots predominantly contained unpalatable sward species, while plots without saplings mainly contained palatable lawn species. After transplantation, sapling survival was higher in exclosures than in the open, and higher in sward exclosures than in lawn exclosures. Sapling growth was higher in swards than in lawns, higher inside than outside exclosures, and higher for Prunus than Crataegus, while browsing on saplings was higher in lawns. Conclusion: Unpalatable swards form essential establishment niches for thorny shrubs in grazed temperate woodlands: they protect against herbivores before thorns fully develop in saplings, and sapling growth is better due to improved micro‐environmental conditions. Once established and thorny, shrub saplings grow out of the protective range of the swards and in turn facilitate tree seedlings, which are essential for long‐term persistence of grazed temperate woodlands. This study shows that nurse plants may start as protégés before becoming facilitators for other plants in a later life stage. This may be common for nurse plants in various ecosystems. We argue that improved understanding of establishment of nurse plants and their constraining factors is crucial for effective conservation and restoration in various ecosystems.     

Differences in pigment composition, photosynthetic rates and chlorophyll fluorescence images of sun and shade leaves of four tree species.

The differential pigment composition and photosynthetic activity of sun and shade leaves of deciduous (Acer pseudoplatanus, Fagus sylvatica, Tilia cordata) and coniferous (Abies alba) trees was comparatively determined by studying the photosynthetic rates via CO(2) measurements and also by imaging the Chl fluorescence decrease ratio (R(Fd)), which is an in vivo indicator of the net CO(2) assimilation rates. The thicker sun leaves and needles in all tree species were characterized by a lower specific leaf area, lower water content, higher total chlorophyll (Chl) a+b and total carotenoid (Cars) content per leaf area unit, as well as higher values for the ratio Chl a/b compared to the much thinner shade leaves and needles that possess a higher Chl a+b and Cars content on a dry matter basis and higher values for the weight ratio Chls/Cars. Sun leaves and needles exhibited higher rates of maximum net photosynthetic CO(2) assimilation (P(Nmax)) measured at saturating irradiance associated with higher maximum stomatal conductance for water vapor efflux. The differences in photosynthetic activity between sun and shade leaves and needles could also be sensed via imaging the Chl fluorescence decrease ratio R(Fd), since it linearly correlated to the P(Nmax) rates at saturating irradiance. Chl fluorescence imaging not only provided the possibility to screen the differences in P(N) rates between sun and shade leaves, but in addition permitted detection and quantification of the large gradients in photosynthetic rates across the leaf area existing in sun and shade leaves.     

Salinity tolerance of 'Valencia' orange trees on rootstocks with contrasting salt tolerance is not improved by moderate shade

The effects of shading in combination with salinity treatments were studied in citrus trees on two rootstocks with contrasting salt tolerance to determine if shading could reduce the negative effects of salinity stress. Well-nourished 2-year-old 'Valencia' orange trees grafted on Cleopatra mandarin (Cleo, relatively salt tolerant) or Carrizo citrange (Carr, relatively salt sensitive), were grown either under a 50% shade cloth or left unshaded in full sunlight. Half the trees received no salinity treatment and half were salinized with 50 mM Cl- during two 9 week salinity periods in the spring and autumn interrupted by an 11 week rainy period. The shade treatment reduced midday leaf temperature and leaf-to-air vapour pressure deficit regardless of salinity treatments. In non-salinized trees, shade increased midday CO2 assimilation rate (A(CO2)) and stomatal conductance, but had no effect on leaf transpiration (E(lf)). Shade also increased leaf chlorophyll and photosynthetic water use efficiency (A(CO2)/E(lf)) in leaves on both rootstocks and increased total plant dry weight in Cleo. The salinity treatment reduced leaf growth and leaf gas exchange parameters. Shade decreased Cl- concentrations in leaves of salinized Carr trees, but had no effect on leaf or root Cl- of trees on Cleo. There were no significant differences in leaf gas exchange parameters of shaded and unshaded salinized plants but the growth reduction from salinity stress was actually greater for shaded than for unshaded trees. Shaded trees on both rootstocks had higher leaf Na+ than unshaded trees after the first salinity period, and this shade-induced elevated leaf Na+ persisted after the second salinity period in trees on Carr. Thus, shading did not alleviate the negative effects of salinity on growth and Na+ accumulation.     

The Effects of Shading and Cutting Treatments on the Photosynthetic Rate of Ryegrass Leaves

As the leaf area of miniature simulated swards of S24 perennialreygrass (Lolium perenne L.) grown in controlled environmentincreased, successive leaves which expanded on the main stemof a sample plant within the sward had progressively lower photosyntheticcapacities. When the plants surrounding the sample plant were cut, the photosyntheticcapacity of the next leaf to expand was increased, whether ornot other leaves on the sample plant had been removed. Whenall fully expanded leaves on the sample plant were removed butthe surrounding plants left uncut, the photosynthetic capacityof successive leaves on the sample plant continued to decreaseas in uncut swards. These results support the view that the changes in photosyntheticcapacity in grass swards are consequences of changes in thelight climate of the developing leaf rather than of changesin demand for assimilates or growth substance supply. The declinein photosynthetic capactiy was contributed to by both stomataland residual diffusion resistances.     

不同生长阶段遮荫对番茄光合作用、干物质分配与叶N、P、K的影响

对番茄植株做了两种不同程度的遮荫处理,观测了夏季午间遮荫对光合速率,干物质积累量及其在根,茎,叶之间的分配,和叶N,P,K的含量以及经济产量的影响,发现不同时期遮荫影响不同。（1）遮荫增加三个阶段（开花早期,盛花期和开花后期）的气孔导度和胞间CO2浓度,显著降低开花早期中午的净化合速率,但盛花期中度遮荫（40%遮荫）使净光合速率随着时间的增加逐渐上升,在开花后期表现更加明显,平均净光合速率比对照高20%以上,蒸腾速率也增加较多。（2）开花早期和盛花期重度遮荫（如本实验中的75%遮荫）显著降低根,茎的干重,而开花后期中度遮荫的根,茎干重高于对照,但遮荫对叶干重的影响不明显。（3）开花早期和盛花期遮荫不明显影响叶片中N,P,K的含量,但开花后期中度遮荫使N,P,K含量增加,（4）开花早期两种遮荫对果实产量影响较小,但盛花期重度遮荫使产量降低,全部产量中无效部分所占的比例上升,开花后期中度遮荫的总产量和有效产量增加,单果重也增加,这些结果表明,在某些时期中度遮荫可以克服夏天辐射过强,气温过高对番茄的不良影响,对番茄生长,干物质积累和提高产量等有利,在生产上有意义。     

Effect of Nitrogen Supply on the Grass and Clover Components of Simulated Mixed Swards Grown under Favourable Environmental Conditions I. Carbon Assimilation and Utilization

Simulated mixed swards of Perennial Ryegrass (Lolium perenneL.) cv. S23 and White clover (Trifolium repens L.) cv. S100were grown from seed under a constant 20 °C day/15 °Cnight temperature regime and their growth and carbon economyexamined. The swards received a nutrient solution daily, whichcontained either High (220 mg l^–1) or Low (10 mg l^–1)nitrate N. Rates of canopy photosynthesis and respiration, and final drymatter yields were similar in the two treatments although theproportions of grass and clover differed greatly. The Low-Nswards were made up largely of clover. The grass plants in theseswards had high root: shoot ratios and low relative photosyntheticrates – both signs of N deficiency – and were clearlyunable to compete with the vigorously growing Low-N clover plants.These had higher relative growth rates and dry matter yieldsthan their High-N counterparts. In the High-N swards clovercontributed around 50 per cent to the sward dry weight throughoutthe measurement period despite having a smaller proportion ofits dry weight in photosynthetic tissue (laminae) than grassover much of it. The latter was compensated for, initially bya higher specific leaf area than grass, and later by a higherphotosynthetic rate per unit leaf weight. The results are discussedin relation to observed declines in the clover content of swardsafter the addition of nitrogen fertilizer in the field. Trifolium repens, white clover, Lolium perenne, perennial ryegrass, nitrogen, photosynthesis, carbon balance     

The functional morphology of juvenile plants tolerant of strong summer drought in shaded forest understories in southern Spain

It has been hypothesized that plants cannot tolerate combined shade and drought, as a result of morphological trade-offs. However, numerous plant species are reportedly widespread in shaded forest understories that face drought, whether seasonal or occasional. We studied juveniles of six plant species that cope with strong summer drought in the understoreys of mixed Quercus forests in southern Spain: the tall-shrubs Phillyrea latifolia and Viburnum tinus, the perennial herb Rubia peregrina, the small shrub Ruscus aculeatus, and climbers Hedera helix and Smilax aspera. All of these species persist in evergreen shade (c. 3% daylight). Two other species were studied as comparators, Ruscus hypoglossum, less tolerant of drought, and Ceratonia siliqua, less tolerant of shade. Morphological and chemical variables relevant to shade and drought tolerance were measured for juveniles in a range of sizes, and also for the leaves of mature plants. The species converge in features that confer tolerance of shade plus drought by reducing demand for resources. Demand for water is reduced through a moderate to high below-ground mass fraction and low to moderate specific leaf area (respectively 0.22–0.52 and 112–172 cm² g^–1 at 1.00 g total dry mass). Demand for both irradiance and water is reduced through a low to moderate foliar nitrogen concentration and long-lived, physically protected leaves (2 yr). The species also converge in features that confer tolerance of either low irradiance or drought through specialized capture of resource, without precluding the other tolerance. These features include deep roots relative to shoot size, moderately higher specific leaf area in shade (1.2–2.0 × that in sun) and higher chlorophyll:nitrogen ratio in shade. Foliar chlorophyll per unit mass was higher in shade, but chlorophyll was not necessarily synthesized in greater amounts; rather, it was higher apparently due to shade effects on structural features linked with specific leaf area. In contrast, N per unit mass was higher in sun leaves independently of specific leaf area. Despite these convergences, the species diverge considerably in their root mass allocation and architecture, leaf saturated water content, density of stomata and guard cell size. No single narrowly defined functional type is needed for tolerance of shade plus drought.     

The photosynthesis of ryegrass leaves grown in a simulated sward

Plants were taken from simulated swards of perennial ryegrass (Lolium perenne) grown in a controlled environment and the rates of photosynthesis of the youngest fully expanded leaves, and the second and third youngest leaves on the same tillers were measured. The youngest leaves had the highest rates and the third the lowest, with the second leaves intermediate. The rate of photosynthesis in bright light of successive youngest expanded leaves decreased as the swards increased in leaf area, but did not when plants were grown so that the main stem was not shaded. When plants were grown at different densities and the photosynthetic rates of leaves of a particular ontogenetic rank were measured, it was found that leaves on plants from higher densities had lower rates of photosynthesis. Also leaves on plants grown in bright light had higher photosynthetic rates than those on plants grown in dim light. It is concluded that the decline in the photosynthetic capacity of successive leaves in a rapidly growing simulated sward is due to the intense shading to which they are subjected during their expansion.