Effects of host shading on consumption and growth of the geometrid Epirrita autumnata: interactive roles of water, primary and secondary compounds

Shading is assumed to reduce allocation to plant phenolics and to defense in general. We here report the results of experimental shading of individual branches or whole canopies in mountain birch on foliar chemistry and on the growth and consumption of a geometrid, Epirritaautumnata. Branch‐wide shading tended to have at least as strong effects on both leaf chemistry and herbivore performance as canopy‐wide shading, indicating local responses of the host to shading. Responses to shading varied among the key leaf traits. Leaf water content was higher and toughness lower in shaded than in non‐shaded leaves. Leaf sugars were lower and protein‐bound and free amino acids higher in shaded than in control leaves. Sucrose and galactose were at high levels in unshaded branches adjacent to shaded ones, suggesting that partial shading enhanced translocation of sugars within canopies. Total phenolics and soluble proanthocyanidins were low in both shading treatments. Of the other phenolic groups, concentrations of gallotannins and cell‐wall‐bound proanthocyanidins did not differ between shaded and non‐shaded leaves. Epirrita larvae grew better in both types of shading treatments compared to either unshaded control trees or to unshaded branches in the branch‐shading trees. By far the most important correlate of larval growth was the amount of water consumed with leaf mass (r=0.94). When variance in water intake was standardized (also largely eliminating parallel variation in proteins), fructose and glucose still had significant positive correlations and proanthocyanidins negative with larval growth on control but not on shade leaves. Concentrations of several phenolic compounds correlated negatively with intake of dry matter and especially water, and different phenolics were important in shaded (gallotannins) and in control (flavonoids) leaves. Our findings strongly suggest that the effects of putatively defensive leaf traits on insect consumption and growth interact with nutritive leaf traits, particularly with water.     

光照对柑橘果皮类胡萝卜素和色泽形成的影响

以“红柿柑”为试材,在柑橘果实膨大末期通过套袋遮光处理以抑制果皮光合作用,研究光照对果皮糖、叶绿素、类胡萝卜素及果实外观色泽的影响。结果表明,遮光后果皮叶绿素含量迅速下降引起果实转色提早,但各种类胡萝卜素含量及总量并未提高,而是显著下降;至果实成熟时由于遮光与光照处理的果皮叶绿素均消失,遮光果实类胡萝卜素含量低颜色变淡,与光照处理相比,遮光前期果皮糖含量下降不大,而后期下降明显;若在后期去袋照光,果皮糖含量上升,与此相应,类胡萝卜素,尤其是β-隐黄质的积累增加,颜色加深,表明光对果皮类胡萝卜素合成尤其是β-隐黄质的积累有促进作用,其原因是光以环境信号的方式影响果皮的类胡萝卜素形成。     

Clonal Plasticity in Response to Partial Neutral Shading in the Stoloniferous Herb Potentilla reptans var. sericophylla

Ramet-pairs of Potentilla reptans L. var.sericophylla Franch from forest gap and forest understory were subjected to unshading, shading and partial shading treatments in a pot experiment. The genet biomass, total length of stolons, number of ramets, specific stolon weight, petiole length and specific petiole weight of the plant species under the shaded condition were smaller than those under the unshaded condition. The stolon internode length did not respond to the various treatments. In the plants from the forest gap, the petiloes of ramet grown in the shaded patch were longer as connected to plant part in the unshaded patch than as connected to plant part under the same shaded condition. Such modification of local response of ramet petiole to shading due to physiological integration was not observed in the plants from the understory. There was no effect of connection to ramets in shaded patches on the local response of the rest ramet characters to the partial unshading.     

匍匐茎草本绢毛匍匐委陵菜对局部遮荫的克隆可塑性

采自林窗和林内生境的绢毛匍匐委陵菜 (PotentillareptansL .var.sericophyllaFranch)“分株对”(即由一匍匐茎节间相连着的两个分株 ,其一为“目标分株” ,另一为“相连分株”)在一户外实验中经历了全不遮荫、全部遮荫和局部遮荫处理。该植物的基株生物量、匍匐茎总长度、分株数、匍匐茎比节间重、叶柄长、比叶柄重在遮荫条件下较小。匍匐茎节间长度没有对遮荫处理发生反应。在局部遮荫处理 ,遮荫斑块的分株的叶柄长度由于连着未遮荫斑块中分株而变得更长。这种克隆整合对克隆形态可塑性的修饰作用只在林窗生境来源的实验植物中观察到。其他克隆生长和克隆形态特征的可塑性在不同生境来源的实验植物间没有差异。     

Shading and the capture of localized soil nutrients: nutrient contents,carbohydrates, and root uptake kinetics of a perennial tussock grass

Summary The ability to exploit spatial and temporal heterogeneity in soil resources can be one factor important to the competitive balance of plants. Competition above-ground may limit selective plant responses to below-ground heterogeneity, since mechanisms such as root proliferation and alterations in uptake kinetics are energy-dependent processes. We studied the effect of shading on the ability of the perennial tussock grassAgropyron desertorum to take up nutrients from enriched soil microsites in two consecutive growing seasons. Roots of unshaded plants selectively increased phosphate uptake capacity in enriched soil microsites (mean increases of up to 73%), but shading eliminated this response. There were no changes in ammonium uptake capacity for roots in control and enriched patches for either shaded or unshaded plants. The 9-day shade treatments significantly reduced total nonstructural carbohydrate (TNC) concentrations for roots in 1990, but had no apparent effect on root carbohydrates in 1991 despite dramatic reductions in shoot TNC and fructan concentrations. Enrichment of the soil patches resulted in significantly greater phosphate concentrations in roots of both shaded and unshaded plants, with less dramatic differences for nitrogen and no changes in potassium concentrations. In many respects the shaded plants did surprisingly well, at least in terms of apparent nutrient acquisition. The effects of aboveground competition on nutrient demand, energy requirements, and belowground processes are discussed for plants exploiting soil resource heterogeneity.     

分株比例对异质光环境下美丽箬竹克隆系统光合生理的影响

分株数量或生物量比例差异会明显影响克隆系统对资源异质性环境的生态适应性, 地下茎木质化、连接稳固的竹类植物在生长过程中相连克隆分株通常会生活在异质光环境中, 但其叶片光合生理特性对异质光环境的响应及其分株比例效应则未见报道。该研究以地下茎相连的美丽箬竹(Indocalamus decorus)克隆系统为实验材料, 设置2个遮光率(分别为50% ± 5%和75% ± 5%)和3个分株比例(遮光与未遮光分株比例分别为1:3、2:2、3:1)处理, 分株数量为4株。分别测定了遮光处理后30、90、150天遮光和未遮光分株叶片光响应特征、气体交换参数、光合色素含量, 分析了异质光环境下美丽箬竹光合生理的变化规律。结果显示: 分株比例对美丽箬竹光合生理有显著影响, 且其与遮光、处理时间交互作用显著。美丽箬竹克隆系统遮光分株比例越大, 即遮光相对分株数量越多, 其表观量子效率(AQE)、光饱和点、最大净光合速率(P_{n max})、净光合速率(P_n)、气孔导度(G_s)、蒸腾速率(T_r)、水分利用效率越大, 光补偿点、暗呼吸速率越小, 其光合效率越高, 光能利用能力越强, 而与之相连的未遮光分株则相反; 随遮光分株比例的增大, 遮光分株叶片叶绿素a、叶绿素b含量呈先升高而后下降的变化趋势, 类胡萝卜素(Car)含量则持续下降, 而与之相连的未遮光分株叶片光合色素含量则呈下降趋势; 遮光率提高, 相同分株比例美丽箬竹克隆系统遮光分株叶片AQE、P_{n max}、P_n、G_s、胞间CO₂浓度(C_i)和光合色素含量总体升高, 而与之相连的未遮光分株叶片P_{n max}、C_i以及Car含量则总体下降。研究结果表明异质光环境下, 遮光分株光合效率和弱光利用能力明显增强, 而未遮光分株则相反, 克隆系统内分株间发生了明显的克隆分工, 且2:2、3:1分株比例克隆系统较1:3分株比例对异质光环境具有更好的适应能力。美丽箬竹克隆系统可通过差异性调节分株光合生理特性和光合色素含量, 提高遮光分株光能利用和光合效率来适应异质光环境, 以提高克隆系统的适合度。     

Why overgrowth of intertidal encrusting algae does not always cause competitive exclusion

Encrusting algae are well-known to be able, for long periods, to withstand shading and overgrowth by other organisms. How this is achieved remains a mystery. It had been proposed that connections with unshaded (non-overgrown) parts of the thallus may allow transfer of nutrients to the shaded part. From this model, I proposed and tested the hypothesis that shaded patches of the intertidal red alga, Hildenbrandia rubra, would survive overgrowth longer, or better, when connected to unshaded thallus than when experimentally separated from surrounding alga. Experimental treatments were shading (black or transparent 80 mm perspex discs or no cover) and scraping (scraped around the disc to remove contacts, a control for effects of scraping, no treatment). The 9 orthogonal combinations of cover and scraping were applied to 3 independent, random replicates (i.e. 27 plots) in each of four randomly chosen sites.

In all 4 sites, over 13 months, shaded H. rubra survived in greater abundance (as % cover) where in contact with surrounding thallus. In one site, there was no effect of shading unless the thallus was isolated. In two sites, shading reduced cover, but was more deleterious where the thallus was isolated. In the fourth site, there were artefacts due to a perspex cover, but still less cover of alga where it was isolated.

This encrusting alga can withstand a long period of complete shading, provided there is connection to unshaded thallus. Interpreting or predicting overgrowth interactions in terms of competitive outcomes is therefore dependent on consideration of whether the overgrown species is actually being affected. It also depends on the duration of overgrowth and, as shown here, the extent to which connectivity with unshaded thallus is effective at preventing or reducing any consequences. Observations and experiments that do not ascertain these are difficult to interpret.     

The Effect of Natural and Artificial Shade on the Macrophytes of Lowland Streams and the Use of Shade as a Management Technique

The reduction of the biomass of three main problem species of submerged aquatic macrophytes was proportional to the light at the stream surface both under artificially shaded sections of stream and in naturally shaded areas when compared to the biomass in adjacent open and unshaded sections of stream. The effect of marginal vegetation in varying the shading effect given to streams of differing width and orientation are described. It is recommended that light should be reduced to about half that presently available in the open, by shading from marginal vegetation but it is warned that too much shade is detrimental to the fish populations of the stream and leads to accentuated local accumulations of leaves. The long term effects are considered but it is expected that partial shading will increase the diversity of submerged plant species. It is suggested that the practice of stream realignment is discontinued and that the natural tendency of streams to create their own meandering channels is allowed but within some defined and generally acceptable framework.     

An effect of shade on the boron requirement for leaf blade elongation in black gram (Vigna mungo L. Hepper)

The present experiment was undertaken to examine whether shading affects the critical boron (B) concentration for leaf blade elongation in black gram. Six days after germination (D6), black gram seedlings were transferred to 8 pairs of pots containing basal nutrient solution: one pot of each pair contained 1 M H₃BO₃ and the other 10 M H₃BO₃. On D10, one day after the emergence of the first trifoliolate leaf blade (TF₁) and one day before the emergence of TF₂, four pairs of pots were shaded, decreasing the light intensity they received in the glasshouse to about 35% of full sunlight compared with 70% received by the unshaded plants. The response to B supply of dry matter (DM) and elongation rate (LBER) of TF₂ were less pronounced in shaded than in unshaded plants. Critical B concentrations for LBER in TF₂ of black gram were 10 mg B kg^-1DM in shaded and 15 mg B unshaded plants, suggesting that shading may have decreased the B requirement for LBER. Thus the present results suggest that light may need to be considered when setting critical values for the diagnosis of B deficiency in black gram.     

Benefits of clonal integration between interconnected ramets of Vallisneria spiralis in heterogeneous light environments

Interconnected ramets of the submersed macrophyte Vallisneria spiralis were subjected to two homogeneous treatments (shading or not shading whole plants) and two heterogeneous treatments (only shading basal or apical ramets of plants). The benefits and costs of clonal integration between connected ramets grown in heterogeneous treatments were examined. Results showed that shading apical ramets induced significant benefits to the performance of whole plant in terms of dry weight per plant (P < 0.01) and number of ramets per plant (P < 0.05). Especially for the unshaded basal ramets, their dry weight, number of ramets, number of branches and total stolon length were 89%, 30%, 29% and 58% higher than the corresponding ramets in homogeneous treatment, respectively. Compared to their controls in homogeneous treatments, unshaded basal ramets produced more leaf mass (0.15 g versus 0.11 g) whereas shaded apical ramets produced more root mass (0.012 g versus 0.008 g). However, there was a different pattern of integration when basal ramets were shaded. Shading basal ramets led to a significant decrease in stolon growth, but the individual performance of shaded ramets improved. Cost-benefit analyses revealed that dry weight per ramet of basal shaded ramets was 31% greater than that of basal shaded ramets in the homogeneous treatment. We can conclude that V. spiralis can benefit from clonal integration in heterogeneous light environments, but that the scale of these benefits is related to the quality of light environments where the clone become established.     

An experimental study of the influence of periphytic algae on invertebrate abundance in a Hong Kong stream

1. Small cages (294cm²) containing unglazed clay quarry tiles were used to investigate the influence of periphytic algae on macroinvertebrate abundance in a Hong Kong stream. Algal biomass was manipulated by shading cages with plastic sheets. Individual cages were assigned to one of three treatment groups: unshaded, shaded and deeply shaded. Invertebrate densities and algal biomass within cages were monitored after 23, 37 and 65 days. 2. Multiple-regression analysis revealed that algal biomass, invertebrate morphospecies richness and total abundance declined with greater shading intensity. The responses of individual invertebrate taxa varied: some (especially Trichoptera) were unaffected by shading, whereas grazers (Baetidae, Psephenidae and Elmidae) declined as shading increased. 3. Significant regressions of the densities of individual taxa upon algal and detrital standing stocks in cages had positive slopes, but algal biomass increased during the study while detrital standing stocks declined. Abundance of invertebrates declined or remained rather stable over time. Density increases resulting from a positive association with algae were apparently offset by declines in abundance correlated with reductions in detritus. 4. Declines in algal biomass were associated with greater shading to which animals may respond directly. To uncouple the link between scarcity of algae and reduction of light intensity, the plastic covers on two groups of cages (deeply shaded and unshaded) which had been placed in the stream for 28 days were reversed so that cages which had been shaded became unshaded and vice versa. The cages were recovered on day 33, Only Coleoptera demonstrated a positive association with atgae inside cages; no relationship between population densities and algal biomass or light intensity was apparent for other taxa. However, the design may have been confounded by deposition of sediment in the cages (due to declining stream discharge) which reduced population densities of colonizers. 5. This study documents changes in invertebrate abundance and morphospecies richness in response periphyton and detritus standing stocks within patches. Summation of such responses may account for observed variations in benthic communities among Hong Kong streams which differ in the extent of shading by riparian vegetation.     

The role of heterotrophic carbon acquisition by the hemiparasitic plant Rhinanthus alectorolophus in seedling establishment in natural communities: a physiological perspective

? Heterotrophic acquisition of substantial amounts of organic carbon by hemiparasitic plants was clearly demonstrated by numerous studies. Many hemiparasites are, however, also limited by competition for light preventing the establishment of their populations on highly productive sites. ? In a growth-chamber experiment, we investigated the effects of competition for light, simulated by shading, on growth and heterotrophic carbon acquisition by the hemiparasite Rhinanthus alectorolophus attached to C(3) and C(4) hosts using analyses of biomass production and stable isotopes of carbon. ? Shading had a detrimental effect on biomass production and vertical growth of the hemiparasites shaded from when they were seedlings, while shading imposed later caused only a moderate decrease of biomass production and had no effect on the height. Moreover, shading increased the proportion of host-derived carbon in hemiparasite biomass (up to 50% in shaded seedlings). ? These results demonstrate that host-derived carbon can play a crucial role in carbon budget of hemiparasites, especially if they grow in a productive environment with intense competition for light. The heterotrophic carbon acquisition can allow hemiparasite establishment in communities of moderate productivity, helping well-attached hemiparasites to escape from the critical seedling stage.     

Interactive effects of shade and surface orientation on the recruitment of spirorbid polychaetes

A series of observations and an experiment were carried out to test hypotheses about the effects of shade on the densities of spirorbid polychaetes (Neodexiospira spp.) on intertidal pneumatophores (mangrove roots) of Avicennia marina. Densities of spirorbids were greater on pneumatophores surrounded by seagrass (Zostera mucronata) than patches without seagrass. Within patches of seagrass, the density and survivorship of spirorbids on pneumatophores was greater near the substratum (covered by seagrass) than high above the substratum (not covered by seagrass). The model that these patterns of abundance are explained by greater recruitment of spirorbids to shaded surfaces was assessed. This was done by experimentally testing the hypothesis that recruitment to patches without seagrass would not differ between the upper (unshaded) and lower surfaces (unshaded) of clear plastic sheets, but would be greater on the lower surfaces (shaded) than upper surfaces (unshaded) of black plastic sheets. Recruitment was consistent with these predictions and therefore provided evidence that differences in densities of spirorbids between substrata with and without seagrass may be due largely to differences in shading.     

Ramet size equalisation in a clonal plant,Phragmites australis

The influence of shading from older generations of dead culms (standing litter) on density, growth rate and development of size structure at the ramet level was investigated in a pure stand of Phragmites australis by experimental neutral shading of plots after removal of standing litter. Initial differences in height distribution between autumn and spring cohorts disappeared in the course of shoot growth. The Gini coefficients of shoot heights and estimated shoot weights indicated that the size structure of the shoots became more equal with increasing mean size in both shaded and unshaded plots. Relative growth rate for height (RHGR) and weight of individual shoots was negatively related to shoot size during the early and presumably storage-dependent growth period, suggesting a strong support for growth of smaller shoots. No etiolation was indicated by mean or maximum height in shaded and unshaded plots, or by the relationship between shoot height and weight. Mean shoot density was significantly lower in shaded than in unshaded plots in one of two shade treatment years. A regression model indicated a small but significant effect of shoot density on the approximately linear relationship between RHGR and the logarithm of height. The growth rate of small shoots was slightly larger at low than at high shoot density. Therefore, it is suggested that the shade from standing litter in P. australis stands can decrease shoot natality in the spring cohort, and thereby increase the support to fewer small shoots.     

Effects of high irradiances on photosynthesis,growth and crassulacean acid metabolism in the epiphyteKalanchoö uniflora

Summary Kalancho? uniflora was grown in the glasshouse with and without shading. Chlorophyll content, area/FW ratio and specific leaf area were higher in leaves of shaded as compared to unshaded plants. Light saturation curves and continuous gas exchange measurements showed that the apparent quantum yield and the light-saturated photosynthetic rate were higher in shaded plants. Shaded plants had lower “mesophyll resistances” than unshaded plants, indicating that the different photosynthetic capacities reflected different contents of ribulose biphosphate carboxylase-oxygenase. Highlight treatment of plants grown in the shade resulted in a decreased photosynthetic efficiency, showing that these plants were sensitive to photoinhibition. However, dry matter production was higher in unshaded than in shaded plants. Obviously the difference in irradiance between the two growth regimes did more than offset the differences in photosynthetic efficiency. Applying additional nutrients did not alter the effects of high PFDs. The results are discussed in respect to photosynthetic performence and plant distribution in the epiphytic habitat.     

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.     

Effects of shading and of seasonal differences in weather on the growth, sugar content and sugar yield of sugar-beet crops

In 1967 and 1968 plots of sugar beet in two identically grown crops were shaded for successive four week periods starting on 13 June, 18 July and 22 August, and the growth of the plants compared with that on unshaded plots. At the beginning of each period in 1967 some shaded and unshaded plants received additional nitrogen, and in 1968 plants continuously shaded from June to September were compared with unshaded plants irrigated to equalize their water losses with those of the shaded plants. The weather in 1967 was sunny and dry and that in 1968 dull and wet, but the yields of dry matter, and particularly of sugar, of the unshaded crop in the 2 years were similar because, although net assimilation rate (E) was greater in 1967 than 1968, mean leaf area index (L) in 1968 was almost double that in the 1967 crop. Shading decreased the incoming radiation by 56%; it decreased E proportionally in 1967, increasing L slightly, but it decreased both E (by 44%) and L in 1968. The weights of dry matter and sugar in the roots of the shaded crop were consistently smaller at the end of shading and at final harvest in October in both years, and their weights, but not those of the tops or the whole plant, at final harvest over all shading treatments in both years were proportional to the amount of radiation received by the crop between June and September. Although shading greatly decreased the supply of photosynthate to the roots, it did not change the sugar content percentage of dry matter, except in the early stages of growth when the sugar content was rapidly increasing. Sugar content percentage of fresh weight of the roots was consistently decreased by shading, wholly because water content was increased relative to dry matter. Therefore the sugar yield of shaded plants was less because the dry weights of the roots were less, not because the partition of photosynthate between sugar storage and root growth changed. There appears to be a mechanism within the root operating over a wide range of photosynthate supply that maintains a nearly constant proportion of sucrose to non-sugar, contrary to the hypothesis that sugar stored in the roots is photosynthate in excess of what can be used in growth of the plant. However, in the extreme condition of continuous shading which drastically decreased the dry weights of all parts of the plant, sugar percentage of dry matter in the roots was decreased, but only from 80 to 70%. In 1967 extra nitrogen applied at the start of shading increased L and the dry weight of the tops in all periods, but had no effect on the dry weight of roots. Because 1968 was a wet year the irrigation treatment had no effect on E or L except for a slight increase in L during the first period; it had no effect on plant dry weight. Both irrigation and additional nitrogen decreased the sugar content percentage of fresh weight of the roots only by altering their water content; sugar percentage of dry matter was unaffected. None of these changes persisted until the final harvest in October.     

Composition of kermes oak browse as affected by shade and stage of maturity

Kermes oak (Quercus coccifera L.) is a sclerophyllous evergreen shrub of the Mediterranean region. In Greece, it grows either alone or in a mixture with other sclerophyllous species in shrublands or as the understory in Pinus brutia forest.The nutritive value and digestibility of kermes oak browse were investigated under 2 light conditions: an an understory species in a P. brutia forest and in adjacent open shrublands. The comparison was made at 3 phenological stages: during the season of rapid growth (in April), after rapid growth was terminated (in May) and when growth had ceased and stems had hardened (in June).Crude protein content of the leaves and twigs was higher in the shaded than in the unshaded plants during the growing season, while the concentrations of total non-structural carbohydrate, cell contents and soluble protein were higher in the unshaded than in the shaded plants. Tannins and lignin content were higher in shaded than in unshaded plants. Similarly, dry matter digestibility was higher in unshaded plants, but declined more drastically in the shaded plants during the growing season. Production was also higher in unshaded than in shaded plants at the end of the growing season.     

Ambient ultraviolet radiation levels in public shade settings

As people become better informed about the harmful effects of prolonged exposure to solar ultraviolet radiation (UVR, 280–400 nm) they will seek the protection of shade, particularly in tropical locations such as Townsville (19° south). Using broad-band radiation sensors for solar ultraviolet-B (280–315 nm), ultraviolet-A (315–400 nm) and daylight (400–800 nm) radiation, the exposure levels were measured in both the horizontal (shaded and unshaded) and vertical (shaded and unshaded) directions. The measurements were conducted at eight locations (shade settings) in Townsville during the period between December 1997 (summer) and May 1998 (beginning of winter). The quality of protection was assessed by the ratio of unshaded to shaded radiation exposure, the UVB/shade protection ratio (UVB-SPR). The UVB-SPR varies considerably between the different shade settings, with a beach umbrella showing the least protection and dense foliage the highest protection. The roof of a house verandah can provide only little protection if the verandah catches the afternoon sun. Increasing cloud cover decreases the UVB-SPR for all settings because of the increase in the diffuse fraction of the radiation. Only one setting provided a UVB-SPR of 15 or higher, as suggested for protective shading against solar UVB radiation. Shade from direct sunlight alone does not provide enough protection against high levels of solar UVR. Apart from the transmission qualities of the shading material, it is the construction of the whole shade setting that determines the exposure levels underneath. A shade structure with enough overhang is recommended so that high levels of scattered radiation do not reach the skin. Received: 26 February 1999 / Accepted: 9 June 1999     

毛乌素沙地旱柳生长和生理特征对遮荫的反应

在一个控制试验中,旱柳经历了全不遮荫、部分遮荫和全部遮荫处理．比较了全不遮荫枝、全部遮荫枝、部分遮荫阳生枝(阳生枝)和部分遮荫阴生枝(阴生枝)的生长和生理特征,结果表明：阳生枝和全部遮荫枝的叶出生率和死亡率分别大于全不遮荫枝和阴生枝;遮荫处理明显影响净光合速率和夜间呼吸速率;阳生枝的分枝生物量、总校长度、枝叶生物量和枝叶重比显著大于全不遮荫枝,而阴生枝的分枝数、分枝生物量、叶面积、叶生物量、基茎、总校长度、枝叶生物量和枝叶重比都显著小于全部遮荫枝．