Aesthetic damage thresholds for twospotted spider mites (Acari: Tetranychidae) on impatiens: effect of plant age and level of infestation

The effects of plant age and infestation level of twospotted spider mite, Tetranychus urticae Koch (Acari: Tetranychidae), on visible plant damage, and the effect of plant age on spider mite population growth on impatiens, Impatiens wallerana Hook.f. (Ericales: Balsaminaceae), were determined by inoculating impatiens plants of three different ages with two densities of spider mites. Each plant was inoculated with either one adult female mite per three leaves or six leaves based on the average number of leaves on plants of each of the three age classes. Subsequently, leaf damage was correlated with mite-days (cumulative spider mite density) per leaf. The results showed that older aged plants exhibited greater damage than younger plants. Regression models of damage thresholds for each plant age suggest that monitoring for spider mites must be done periodically throughout the entire plant production cycle, but that more attention should be given toward the end of the cycle. Measurements of visible leaf damage were correlated with plant marketability. Specifically, the level of damage (proportion of damaged leaves per plant) at which plant marketability changes from a "premium" to a "discounted" category was 0.04-0.06. Thus, regression equations of the damage threshold could be used to estimate a cumulative spider mite density or mite-days equivalent to the economic threshold. Based on these equations, 5% leaf damage corresponds to 2.1, 1.51, and 1.25 mite-days for youngest, intermediate, and oldest plants, respectively. Because the damage threshold on impatiens was shown to be very low, the action threshold for biological control is essentially zero, and predators would need to be released as soon as damage is observed.     

Impact of twospotted spider mite (Acari: Tetranychidae) on growth and productivity of glasshouse cucumbers

Experiments were conducted to assess the damage of the glasshouse cucumber by twospotted spider mite, Tetranychus urticae Koch, and to investigate when the economic yield begins to decrease after T. urticae infestations. To assess the damage, dry matter partitioning in the cucumber plant was quantified and plant growth analyses were conducted at four different T. urticae infestation levels. T. urticae infestations decreased leaf productivity by reducing the total number of leaves per plant. Approximately 14% reductions of total leaf areas could result in significant yield loss. The decreased leaf productivity by T. urticae feeding caused biomass reductions and altered the pattern of dry matter partitioning in the plant; damaged plants accumulated more dry matter in the leaf, and partitioning of dry matter to fruits was hindered. The economic yield of cucumber began to significantly decrease as early as 4 wk after heavy mite infestations. This study also showed the seasonal differences in T. urticae-cucumber damage interactions among mite infestation levels.     

Seasonal dynamics of economic injury levels for Tetranychus urticae Koch (Acari, Tetranychidae) on Cucumis sativus L.

Abstract:  Economic injury level (EIL) for Tetranychus urticae Koch on glasshouse cucumbers was established in two ways: EILs based on the number of T. urticae introduced to cucumber early in the growing season and EILs based on mite density and feeding durations (i.e. cumulative mite-days) in each of four growing seasons. The relationship between yield loss and T. urticae infestation level varied among four growing seasons. One mite introduced to cucumber early in the growing season could reduce the cucumber yield by 5.03, 3.20, 1.12 and 2.86 g per plant in spring, summer, autumn and winter, respectively. One mite-day (i.e. one mite feeding in a day) reduced cucumber yield by 0.0123, 0.0074, 0.0035 and 0.0073 g per plant in spring, summer, autumn and winter, respectively. When the seasonal dynamics of market values of cucumber was considered, EILs were the lowest in spring and the highest in autumn. When four seasonal data were combined, EILs can be calculated as EIL = C /(0.00276 ×  K ) for the number of T. urticae introduced early in the growing season and EIL = C /(6.8342 × 10⁻⁶ ×  K ) for cumulative mite-days, where K is efficacy of control measure and C the management cost. This study demonstrated the seasonal dynamics of EILs for T. urticae on glasshouse cucumbers, suggesting that decision making for T. urticae management on cucumber needs to be seasonally different under the glasshouse condition.     

Spatial distribution of phytophagous mites (Acari: Tetranychidae) on strawberry plants

Many phytophagous mites can attack strawberry plants, Fragaria x ananassa, among them the southern red mite, Oligonychus ilicis McGregor, and the two-spotted spider mite, Tetranychus urticae Koch. They are found together feeding on the same plant on the upper and underside of the leaves, respectively. Here we studied the choice for feeding sites of O. ilicis and T. urticae on strawberry plants. The first hypothesis tested whether the feeding site choice would be related to the fitness of the species. The second hypothesis dealt whether the feeding site would be determined by the presence of a heterospecific mite. We evaluated the preference, biology and reproductive success of O. ilicis and T. urticae on the under and upper side surface of strawberry leaves infested or not by the heterospecific. O. ilicis preferred to stay on the upper side surface while T. urticae preferred the underside. The preference for the leaf surface correlated with the reproductive success of the species (measured by the intrinsic growth rate). The choice pattern of feeding sites did not alter when the choice test was applied using sites previously infested by heterospecific. Although O. ilicis and T. urticae, apparently, do not interact directly for feeding sites, there is a chance that the first species induces defenses in strawberry plant enabling to reduce the fitness of the second species. The possibility of those species stay together on strawberry plant increases the damage capacity to the culture.     

A herbivorous mite down-regulates plant defence and produces web to exclude competitors

Herbivores may interact with each other through resource competition, but also through their impact on plant defence. We recently found that the spider mite Tetranychus evansi down-regulates plant defences in tomato plants, resulting in higher rates of oviposition and population growth on previously attacked than on unattacked leaves. The danger of such down-regulation is that attacked plants could become a more profitable resource for heterospecific competitors, such as the two-spotted spider mite Tetranychus urticae. Indeed, T. urticae had an almost 2-fold higher rate of oviposition on leaf discs on which T. evansi had fed previously. In contrast, induction of direct plant defences by T. urticae resulted in decreased oviposition by T. evansi. Hence, both herbivores affect each other through induced plant responses. However, when populations of T. evansi and T. urticae competed on the same plants, populations of the latter invariably went extinct, whereas T. evansi was not significantly affected by the presence of its competitor. This suggests that T. evansi can somehow prevent its competitor from benefiting from the down-regulated plant defence, perhaps by covering it with a profuse web. Indeed, we found that T. urticae had difficulties reaching the leaf surface to feed when the leaf was covered with web produced by T. evansi. Furthermore, T. evansi produced more web when exposed to damage or other cues associated with T. urticae. We suggest that the silken web produced by T. evansi serves to prevent competitors from profiting from down-regulated plant defences.     

Phytoseiulus persimilis response to herbivore-induced plant volatiles as a function of mite-days

The predatory mite, Phytoseiulus persimilis (Acari: Phytoseiidae), uses plant volatiles (i.e., airborne chemicals) triggered by feeding of their herbivorous prey, Tetranychus urticae (Acari: Tetranychidae), to help locate prey patches. The olfactory response of P. persimilis to prey-infested plants varies in direct relation to the population growth pattern of T. urticae on the plant; P. persimilis responds to plants until the spider mite population feeding on a plant collapses, after which infested plants do not attract predators. It has been suggested that this represents an early enemy-free period for T. urticae before the next generation of females is produced. We hypothesize that the mechanism behind the diminished response of predators is due to extensive leaf damage caused by T. urticae feeding, which reduces the production of volatiles irrespective of the collapse of T. urticae population on the plant. To test this hypothesis we investigated how the response of P. persimilis to prey-infested plants is affected by: 1) initial density of T. urticae, 2) duration of infestation, and 3) corresponding leaf damage due to T. urticae feeding. Specifically, we assessed the response of P. persimilis to plants infested with two T. urticae densities (20 or 40 per plant) after 2, 4, 6, 8, 10, 12 or 14 days. We also measured leaf damage on these plants. We found that predator response to T. urticae-infested plants can be quantified as a function of mite-days, which is a cumulative measure of the standing adult female mite population sampled and summed over time. That is, response to volatiles increased with increasing numbers of T. urticae per plant or with the length of time plant was infested by T. urticae, at least as long at the leaves were green. Predatory mites were significantly attracted to plants that were infested for 2 days with only 20 spider mites. This suggests that the enemy-free period might only provide a limited window of opportunity for T. urticae because relatively low numbers of T. urticae per plant can attract predators. Leaf damage also increased as a function of mite-days until the entire leaf was blanched. T. urticae populations decreased at this time, but predator response to volatiles dropped before the entire leaf was blanched and before the T. urticae population decreased. This result supports our hypothesis that predator response to plant volatiles is linked to and limited by the degree of leaf damage, and that the quantitative response to T. urticae populations occurs only within a range when plant quality has not been severely compromised.     

Gene Pyramiding of Peptidase Inhibitors Enhances Plant Resistance to the Spider Mite Tetranychus urticae

The two-spotted spider mite Tetranychus urticae is a damaging pest worldwide with a wide range of host plants and an extreme record of pesticide resistance. Recently, the complete T. urticae genome has been published and showed a proliferation of gene families associated with digestion and detoxification of plant secondary compounds which supports its polyphagous behaviour. To overcome spider mite adaptability a gene pyramiding approach has been developed by co-expressing two barley proteases inhibitors, the cystatin Icy6 and the trypsin inhibitor Itr1 genes in Arabidopsis plants by Agrobacterium-mediated transformation. The presence and expression of both transgenes was studied by conventional and quantitative real time RT-PCR assays and by indirect ELISA assays. The inhibitory activity of cystatin and trypsin inhibitor was in vitro analysed using specific substrates. Single and double transformants were used to assess the effects of spider mite infestation. Double transformed lines showed the lowest damaged leaf area in comparison to single transformants and non-transformed controls and different accumulation of H(2)O(2) as defence response in the leaf feeding site, detected by diaminobenzidine staining. Additionally, an impact on endogenous mite cathepsin B- and L-like activities was observed after feeding on Arabidopsis lines, which correlates with a significant increase in the mortality of mites fed on transformed plants. These effects were analysed in view of the expression levels of the target mite protease genes, C1A cysteine peptidase and S1 serine peptidase, identified in the four developmental mite stages (embryo, larvae, nymphs and adults) performed using the RNA-seq information available at the BOGAS T. urticae database. The potential of pyramiding different classes of plant protease inhibitors to prevent plant damage caused by mites as a new tool to prevent pest resistance and to improve pest control is discussed.     

叶片物理性状对玉米杂交种京科968防御二斑叶螨危害的影响

叶螨(Acari:Tetranychidae)危害是造成玉米减产的重要原因之一,其中二斑叶螨Tetranychus urticae Koch是我国玉米Zea mays L.生产中的主要害螨之一.抗螨玉米品种的选育是有效防治叶螨的途径之一.本研究以我国广泛种植的玉米杂交种京科968及其母本京724、父本京92,先玉335...     

Photosynthetic Carbon Metabolism in the Palisade Parenchyma and Spongy Parenchyma of Vicia faba L

Palisade parenchyma cells and spongy parenchyma cells were isolated separately from Vicia faba L. leaflets. Extracts of the cell isolates were assayed for several enzymes involved in CO₂ fixation and photorespiration. When compared on a chlorophyll basis, the levels of enzyme activities either were equal in the different cell types or were greater in the spongy parenchyma; this difference is a reflection, perhaps, of the higher protein-chlorophyll ratio in the latter tissue. The distribution of radioactivity in the products of photosynthesis by each cell type was the same at various times after exposure to NaH¹⁴CO₃, and the kinetics of ¹⁴C incorporation into these compounds was similar. However, a larger percentage of radioactivity was incorporated by the cell isolates into the 80% ethanol-insoluble fraction and correspondingly less into the neutral fraction as compared to whole leaf. It was concluded that photosynthetic CO₂ fixation is similar in the different mesophyll tissues from which these cells were derived.     

Damage on 'Hass' avocado leaves, webbing and nesting behaviour of Oligonychus perseae (Acari: Tetranychidae)

The damage of Oligonychus perseae Tuttle, Baker and Abbatiello on 'Hass' avocado trees occurs mainly on the underside of the leaves along the midrib, main veins and leaf depressions. The lower epidermal, spongy parenchyma and palisade parenchyma cells of the leaf tissues are destroyed. Large necrotic areas on the underside of the leaves result from feeding when high population levels occur. Feeding and reproduction takes place in 'nests' of silken webbing, which also provide protection from some predator mites and other natural enemies. Oligonychus perseae shows a modification of the earlier defined life-type web nest (WN-c). The greatest number of nests built by a female was 12.17 at 20°C and the greatest number of eggs per female per nest was 5.20 at 25°C.     

Acaricidal properties of spinosad against Tetranychus urticae and Panonychus ulmi (Acari: Tetranychidae)

Laboratory bioassays were conducted to characterize the activity of the insecticide spinosad against the twospotted spider mite, Tetranychus urticae Koch, and European red mite, Panonychus ulmi (Koch) (Acari: Tetranychidae). T. urticae females and larvae were individually placed on bean, Phaseolus vulgaris L. (Fabaceae), leaf disks treated with four rates of spinosad (25, 55, 121, and 266 ppm) and a water control. Significantly fewer T. urticae completed development on any spinosad rates (<15%) compared with the control (>85%), whereas spinosad exhibited no significant effects on P. ulmi development; 72.5 and 83.1% of P. ulmi completed development on apple (Malus pumila P. Mill, Rosaceae) leaf disks treated with 75 ppm spinosad and the control, respectively. T. urticae adult females placed on spinosad-treated disks had significantly higher mortality and lower oviposition rates compared with the water control; no significant mortality effects were observed until 3 d after placing adults on leaf disks. In choice tests where half of a bean leaf was treated with 55 ppm spinosad transversally or longitudinally, T. urticae females were repelled by spinosad and largely oviposited and fed on nonspinosad treated areas. Spinosad did not affect the behavior of P. ulmi females. When T. urticae females were released on potted bean plants (two-leaf stage) in which leaves received spinosad sprays on the adaxial or abaxial leaf surfaces, or complete spinosad coverage on one or two of the leaves, mite population increase lagged significantly behind those released on control plants. These results indicate that spinosad has significant acaricidal effects against T. urticae but not P. ulmi.     

Effect of different densities of the twospotted spider mite Tetranychus urticae on CO2 assimilation, transpiration, and stomatal behaviour in rose leaves

The effect of population density of Tetranychus urticae Koch on CO2 assimilation, transpiration and stomatal behaviour in rose leaves and on the diameter and length of stems and flower buds was investigated under greenhouse conditions. The investigation was performed in order to gain more insight into integrated control systems in rose crops grown under greenhouse conditions. Physiological processes, such as photosynthesis and transpiration, as well as stomatal behaviour and chlorophyll content, were studied as they form part of the plant's nutrition mechanism and therefore affect the quantity and quality of the flowers. Information related to the effect of spider mite population density on bloom quality, diameter and length of stems and flower buds was also collected. The data indicate that increased mite density coincides with a decrease in the net photosynthetic rate, transpiration and chlorophyll content. Higher mite densities on leaves cause stomata to remain open for longer periods, which allows a greater loss of water. Spider mite densities of 10 and 50 mites per leaf cause a reduction in flower stem length of 17 and 26%, respectively, as compared to plants with no mites present.     

Allocating time between feeding, resting and moving by the two-spotted spider mite, Tetranychus urticae and its predator Phytoseiulus persimilis

This study characterizes the timing of feeding, moving and resting for the two-spotted spider mite, Tetranychus urticae Koch and a phytoseiid predator, Phytoseiulus persimilis Athias-Henriot. Feeding is the interaction between T. urticae and plants, and between P. persimilis and T. urticae. Movement plays a key role in locating new food resources. Both activities are closely related to survival and reproduction. We measured the time allocated to these behaviours at four ages of the spider mite (juveniles, adult females immediately after moult and adult females 1 and 3 days after moult) and two ages of the predatory mite (juveniles and adult females). We also examined the effect of previous spider mite-inflicted leaf damage on the spider mite behaviour. Juveniles of both the spider mite and the predatory mite moved around less than their adult counterparts. Newly emerged adult female spider mites spent most of their time moving, stopping only to feed. This represents the teneral phase, during which adult female spider mites are most likely to disperse. With the exception of this age group, spider mites moved more and fed less on previously damaged than on clean leaves. Because of this, the spider mite behaviour was initially more variable on damaged leaves. Phytoseiulus persimilis rested at all stages for a much larger percentage of the time and spent less time feeding than did T. urticae; the predators invariably rested in close proximity to the prey. Compared to adult predators, juveniles spent approximately four times as long handling a prey egg. The predator-prey interaction is dependent upon the local movement of both the predators and prey. These details of individual behaviours in a multispecies environment can provide an understanding of population dynamics.     

Ontogenetic differences in mesophyll structure and chlorophyll distribution in Eucalyptus globulus ssp. globulus

Mesophyll structure has been associated with the photosynthetic performance of leaves via the regulation of internal light and CO(2) profiles. Differences in mesophyll structure and chlorophyll distribution within three ontogenetically different leaf types of Eucalyptus globulus ssp. globulus were investigated. Juvenile leaves are blue-grey in color, dorsiventral (adaxial palisade layer only), hypostomatous, and approximately horizontal in orientation. In contrast, adult leaves are dark green in color, isobilateral (adaxial and abaxial palisade), amphistomatous, and nearly vertical in orientation. The transitional leaf type has structural features that appear intermediate between the juvenile and adult leaves. The ratio of mesophyll cell surface area per unit leaf surface area (A(mes)/A) of juvenile leaves was maximum at the base of a single, adaxial palisade layer and declined through the spongy mesophyll. Chlorophyll a + b content showed a coincident pattern, while the chlorophyll a:b ratio declined linearly from the adaxial to abaxial epidermis. In comparison, the mesophyll of adult leaves had a bimodal distribution of A(mes)/A, with maxima occurring beneath both the adaxial and abaxial surfaces within the first layer of multiple palisade layers. The distribution of chlorophyll a + b content had a similar pattern, although the maximum ratio of chlorophyll a:b occurred immediately beneath the adaxial and abaxial epidermis. The matching distributions of A(mes)/A and chlorophyll provide further evidence that mesophyll structure may act to influence photosynthetic performance. These changes in internal leaf structure at different life stages of E. globulus may be an adaptation for increased xeromorphy under increasing light exposure experienced from the seedling to adult tree, similar to the characteristics reported for different species according to sunlight exposure and water availability within their native habitats.     

Interactions in a tritrophic acarine predator-prey metapopulation system III: effects of Tetranychus urticae (Acari: Tetranychidae) on host plant condition

Spider mites are serious pests on many economically important plant species, because they may reduce plant productivity and, at high mite densities, overexploit and even kill the host plants. We have conducted a series of greenhouse experiments to quantify the effects of two-spotted spider mites (Tetranychus urticae) on host plants (Phaseolusvulgaris). The average amount of chlorophyll per cm² leaf area was used as a measure of plant condition. It was shown that chlorophyll concentration decreases with plant age and intensity of spider mite feeding. Damage caused by spider mites was assessed visually, using the Leaf Damage Index (LDI) defined by, and a mathematical relationship between the visual measurements and the amount of chlorophyll/cm² was fitted to data. The relationship may serve as a short-cut to estimate overall plant injury, expressed as the relative loss of chlorophyll/cm² leaf area caused by spider mites (D). D takes values between 0 (no injury) and 1 (all leaves dead). A highly significant positive relationship between the instantaneous spider mite density and D was found, even though D is expected to reflect the cumulated density of mites (mite-days). A model of plant growth incorporating information about plant age and D predicts that plant area has a maximum when plant age is about 60 days, and that plant area decreases exponentially with an increase in D. This revised version was published online in July 2006 with corrections to the Cover Date.     

Within-plant distribution of twospotted spider mite, Tetranychus urticae Koch (Acari: Tetranychidae), on ivy geranium: development of a presence-absence sampling plan

The twospotted spider mite, Tetranychus urticae Koch, is an important pest of ivy geranium and other ornamental plants. As a part of our long-term goal to develop an integrated crop management program for ivy geraniums, the focus of this study was to produce a reliable sampling method for T. urticae on this bedding plant. Within-plant mite distribution data from a greenhouse experiment were used to identify the young-fully-opened leaf as the sampling unit. We found that 53% of the mites on a plant are on the young-fully-opened leaves. On average 22, 37, and 41% of the leaves belonged to the young, young-fully-opened, and old leaf categories, respectively. We then developed a presence-absence sampling method for T. urticae in ivy geranium using generic Taylor's coefficients for this pest. We found the optimal binomial sample sizes for estimating populations of T. urticae at densities of between 0 and 3 mites/leaf to be quite large; therefore, we recommend the use of numerical sampling within this range of T. urticae densities. We also suggest that population estimates of T. urticae on ivy geranium be done based on mite density/unit area of greenhouse space, both for conventional greenhouse pest management, and for determining how many phytoseid predators to release when using biological control.     

Disease prevalence and transmission of Microsporidium phytoseiuli infecting the predatory mite, Phytoseiulus persimilis (Acari: Phytoseiidae)

Isolated colonies of the predatory mite, Phytoseiulus persimilis, were used to gain information regarding prevalence and transmission of Microsporidium phytoseiuli. Two colonies of P. persimilis were reared on spider mite (Tetranychus urticae)-infested bean plants in isolated cages. Disease prevalence of predators from Colony 1 remained relatively low (between 0 and 15%) over 57 weeks of observation whereas disease prevalence of predators from Colony 2 increased over 3 months (from 12 to 100%). Disease prevalence among predators from Colony 1 had increased to 100% 2 months after weekly sampling had ceased for this colony and periodic sampling confirmed that disease prevalence among individuals of both colonies remained at 100%. Microsporidian spores were not detected in randomly chosen samples of T. urticae prey mites that were removed and examined biweekly during this period. Although numerous microsporidian spores were observed in smear preparations of fecal pellets examined by light microscopy, spores were not observed on leaf surfaces or predator feces when examined by SEM. The latter appeared as intact aggregates composed of numerous dumbbell-shaped crystals and it is unlikely that spores are liberated from intact fecal pellets onto leaf surfaces. Vertical transmission of M. phytoseiuli was 100%; horizontal transmission was low (14.3%) and occurred only when immature P. persimilis were permitted to develop in contact with infected immature and adult predators. The mean number of eggs produced per mated pair was highest when uninfected females were mated with uninfected males (63.2 eggs per mated pair). Although mean egg production decreased when one or both parents were infected, not all differences were significant. Male predatory mites did not contribute to infection of their progeny. Results suggest that routine examination of P. persimilis for microsporidian spores is essential for the management of M. phytoseiuli within P. persimilis colonies. Low disease prevalence and lack of obvious disease signs or symptoms, as in the case of M. phytoseiuli, increase the probability that these pathogens will escape notice unless individuals are routinely examined for pathogens.     

叶螨及两种替代食物对尼氏真绥螨发育和繁殖的影响

在温度25 ℃、相对湿度80％条件下,以油茶花粉、二斑叶螨和腐食酪螨3种食物饲喂尼氏真绥螨,研究不同食物对尼氏真绥螨生长发育和繁殖的影响.结果表明：尼氏真绥螨虽能取食腐食酪螨的卵,但因摄食量不足导致不能完成正常发育;而取食油茶花粉和二斑叶螨能正常完成发育,并进行繁殖.取食油茶花粉的尼氏真绥螨全世代发育历期、雌螨寿命、产卵历期、平均每雌总产卵量分别为6.18 d、24.97 d、16.72 d和23.03粒,取食二斑叶螨的尼氏真绥螨分别为5.67 d、25.72 d、18.17 d和25.38粒;以二斑叶螨为食的尼氏真绥螨发育快,寿命和产卵历期长,平均每雌产卵量高.以油茶花粉和二斑叶螨为食的尼氏真绥螨实验种群数量动态均呈上升趋势,其中以油茶花粉饲养的种群趋势指数最高（I=14.28）,以二斑叶螨饲养的种群加倍时间最短（t=3.5201 d）.     

Photosynthetic and structural acclimation to light direction in vertical leaves of Silphium terebinthinaceum

The azimuth of vertical leaves of Silphium terebinthinaceum profoundly influenced total daily irradiance as well as the proportion of direct versus diffuse light incident on the adaxial and abaxial leaf surface. These differences caused structural and physiological adjustments in leaves that affected photosynthetic performance. Leaves with the adaxial surface facing East received equal daily integrated irradiance on each surface, and these leaves had similar photosynthetic rates when irradiated on either the adaxial or abaxial surface. The adaxial surface of East-facing leaves was also the only surface to receive more direct than diffuse irradiance and this was the only leaf side which had a clearly defined columnar palisade layer. A potential cost of constructing East-facing leaves with symmetrical photosynthetic capcity was a 25% higher specific leaf mass and increased leaf thickness in comparison to asymmetrical South-facing leaves. The adaxial surface of South-facing leaves received approximately three times more daily integrated irradiance than the abaxial surface. When measured at saturating CO₂ and irradiance, these leaves had 42% higher photosynthetic rates when irradiated on the adaxial surface than when irradiated on the abaxial surface. However, there was no difference in photosynthesis for these leaves when irradiated on either surface when measurements were made at ambient CO₂. Stomatal distribution (mean adaxial/abaxial stomatal density = 0.61) was unaffected by leaf orientation. Thus, the potential for high photosynthetic rates of adaxial palisade cells in South-facing leaves at ambient CO₂ concentrations may have been constrained by stomatal limitations to gas exchange. The distribution of soluble protein and chlorophyll within leaves suggests that palisade and spongy mesophyll cells acclimated to their local light environment. The protein/chlorophyll ratio was high in the palisade layers and decreased in the spongy mesophyll cells, presumably corresponding to the attentuation of light as it penetrates leaves. Unlike some species, the chlorophyll a/b ratio and the degree of thylakoid stacking was uniform throughout the thickness of the leaf. It appears that sun-shade acclimation among cell layers of Silphium terebinthinaceum leaves is accomplished without adjustment to the chlorophyll a/b ratio or to thylakoid membrane structure.     

二斑叶螨为害对草莓叶片H2O2、MDA含量以及部分防御酶活性的影响

本文旨在探究二斑叶螨Tetranychus urticae为害对草莓Fragaria×ananassa Duch.叶片内过氧化氢（H2O2）、丙二醛（MDA）含量以及部分防御酶活性的影响。在草莓苗上接种不同数量（5~25头）的二斑叶螨,分别在接种后的24 h、48 h和72 h取样,分析草莓叶片内H2O2、MDA的含量以及部分防御酶的活性。结果显示,二斑叶螨为害的草莓叶片内H2O2和MDA的含量以及超氧化物歧化酶（SOD）的活性随着时间的延长而呈现先升后降的趋势,在二斑叶螨持续为害草莓叶片24 h、48 h和72 h时,受损草莓叶片中H2O2的含量均显著高于对照（P<0.05）,不同密度二斑叶螨为害的草莓叶片中H2O2的含量均显著高于对照（P<0.05）,但与取食时间关系不大。当为害时间达到48 h时,MDA的含量和SOD的活性均达到最高峰,此时它们均与二斑叶螨的密度密切相关。当二斑叶螨为25头/叶时,MDA的含量和SOD的活性分别约是对照的3.6倍和10倍。过氧化物酶（POD）和过氧化氢酶（CAT）的活性随时间延长不断升高,均在二斑叶螨为害72 h时达到最高峰。同时,二斑叶螨的为害时间和为害密度之间存在一定的交互作用。以上结果表明草莓叶片主要通过调节其体内H2O2和MDA的含量以及各种防御酶活性的变化,对二斑叶螨的为害产生应激反应。