Solar ultraviolet-B radiation affects plant-insect interactions in a natural ecosystem of Tierra del Fuego (southern Argentina)

We examined the effects of solar ultraviolet-B radiation (UVB) on plant-herbivore interactions in native ecosystems of the Tierra del Fuego National Park (southern Argentina), an area of the globe that is frequently under the Antarctic “ozone hole” in early spring. We found that filtering out solar UVB from the sunlight received by naturally-occurring plants of Gunnera magellanica, a creeping perennial herb, significantly increased the number of leaf lesions caused by chewing insects. Field surveys suggested that early-season herbivory was principally due to the activity of moth larvae (Lepidoptera: Noctuidae). Manipulative field experiments showed that exposure to solar UVB changes the attractiveness of G. magellanica leaf tissue to natural grazers. In a laboratory experiment, locally caught moth caterpillars tended to eat more tissue from leaves grown without UVB than from leaves exposed to natural UVB during development; however, the difference between treatments was not significant. Leaves grown under solar UVB had slightly higher N levels than leaves not exposed to UVB; no differences between UVB treatments in specific leaf mass, relative water content, and total methanol-soluble phenolics were detected. Our results show that insect herbivory in a natural ecosystem is influenced by solar UVB, and that this influence could not be predicted from crude measurements of leaf physical and chemical characteristics and a common laboratory bioassay. Received: 4 December 1997 / Accepted: 12 May 1998     

Genetic variation in defensive chemistry in Plantago lanceolata (Plantaginaceae) and its effect on the specialist herbivore Junonia coenia (Nymphalidae)

To examine genetic variation in defensive chemistry within and between natural populations of Plantago lanceolata, we performed a greenhouse experiment using clonal replicates of 15 genotypes from each of two populations, from a mowed lawn and an abandoned hayfield. Replicates of each genotype were harvested for determinations of aboveground biomass and leaf chemical content either at the beginning of the experiment (initial controls), after exposure to herbivory by larvae of Junonia coenia, a specialist on P. lanceolata (herbivory treatment), or at the end of the experiment without exposure to herbivory (final controls). Allocation to the iridoid glycosides aucubin and catalpol and the phenylpropanoid glycoside verbascoside displayed significant genetic variation within and between populations, and differed with leaf age. Significant genotypextreatment interactions indicated genetic variation in response of leaf chemistry to the treatments. There was no evidence for a cost of allocation to chemical defense: genetic correlations within and between chemical pathways and between defensive chemicals and aboveground growth were positive or nonsignificant. Although iridoid glycosides are known to be qualitative feeding stimulants for J. coenia, multiple regression of larval survivorship on leaf chemical content and shoot biomass indicated that larvae had lower survivorship on P. lanceolata ge-notypes with higher concentrations of aucubin in the leaves. Larval survivorship was unaffected by levels of catalpol and verbascoside. Thus, although specialist herbivores may respond to defensive chemicals as qualitative feeding stimulants, they do not necessarily have higher fitness on plant genotypes containing higher concentrations of these chemicals.     

Leaf Colour as a Signal of Chemical Defence to Insect Herbivores in Wild Cabbage (Brassica oleracea)

Leaf colour has been proposed to signal levels of host defence to insect herbivores, but we lack data on herbivory, leaf colour and levels of defence for wild host populations necessary to test this hypothesis. Such a test requires measurements of leaf spectra as they would be sensed by herbivore visual systems, as well as simultaneous measurements of chemical defences and herbivore responses to leaf colour in natural host-herbivore populations. In a large-scale field survey of wild cabbage (Brassica oleracea) populations, we show that variation in leaf colour and brightness, measured according to herbivore spectral sensitivities, predicts both levels of chemical defences (glucosinolates) and abundance of specialist lepidopteran (Pieris rapae) and hemipteran (Brevicoryne brassicae) herbivores. In subsequent experiments, P. rapae larvae achieved faster growth and greater pupal mass when feeding on plants with bluer leaves, which contained lower levels of aliphatic glucosinolates. Glucosinolate-mediated effects on larval performance may thus contribute to the association between P. rapae herbivory and leaf colour observed in the field. However, preference tests found no evidence that adult butterflies selected host plants based on leaf coloration. In the field, B. brassicae abundance varied with leaf brightness but greenhouse experiments were unable to identify any effects of brightness on aphid preference or performance. Our findings suggest that although leaf colour reflects both levels of host defences and herbivore abundance in the field, the ability of herbivores to respond to colour signals may be limited, even in species where performance is correlated with leaf colour.     

DNA barcoding of the endemic New Zealand leafroller moth genera, Ctenopseustis and Planotortrix

Molecular techniques such as DNA barcoding have become popular in assisting species identification especially for cryptic species complexes. We have analysed data from a 468-bp region of the mitochondrial cytochrome oxidase subunit I (COI) gene from 200 specimens of 12 species of endemic New Zealand leafroller moths (Tortricidae) from the genera Planotortrix and Ctenopseustis to assess whether the DNA barcoding region can distinguish these species. Among the 200 sequences analysed, 72 haplotypes were recovered, with each genus forming a separate major clade. Maximum likelihood phylogenetic methods were used to test whether species fell into reciprocally monophyletic clades. The optimal phylogeny showed that four species within the genus Ctenopseustis (C. obliquana, C. herana, C. filicis and C. fraterna) and three within Planotortrix (P. octo, P. excessana and P. avicenniae) are polyphyletic. Shimodaira-Hasegawa tests rejected a null hypothesis of monophyly for the species C. obliquana, C. herana, P. octo and P. excessana. Comparisons of within and between species levels of sequence divergence for the same set of seven species showed cases where maximum levels of within-species divergence were greater than some levels of between-species divergence. DNA barcoding using this region of the COI gene is able to distinguish the two genera and some species within each genus; however, many species cannot be identified using this method. Finally, we discuss the possible reasons for this polyphyly, including incomplete lineage sorting, introgression, horizontal gene transfer and incorrect taxonomy.     

The Only African Wild Tobacco,Nicotiana africana: Alkaloid Content and the Effect of Herbivory

Herbivory in some Nicotiana species is known to induce alkaloid production. This study examined herbivore-induced defenses in the nornicotine-rich African tobacco N. africana, the only Nicotiana species indigenous to Africa. We tested the predictions that: 1) N. africana will have high constitutive levels of leaf, flower and nectar alkaloids; 2) leaf herbivory by the African bollworm Helicoverpa armigera will induce increased alkaloid levels in leaves, flowers and nectar; and 3) increased alkaloid concentrations in herbivore-damaged plants will negatively affect larval growth. We grew N. africana in large pots in a greenhouse and exposed flowering plants to densities of one, three and six fourth-instar larvae of H. armigera, for four days. Leaves, flowers and nectar were analyzed for nicotine, nornicotine and anabasine. The principal leaf alkaloid was nornicotine (mean: 28 µg/g dry mass) followed by anabasine (4.9 µg/g) and nicotine (0.6 µg/g). Nornicotine was found in low quantities in the flowers, but no nicotine or anabasine were recorded. The nectar contained none of the alkaloids measured. Larval growth was reduced when leaves of flowering plants were exposed to six larvae. As predicted by the optimal defense theory, herbivory had a localized effect and caused an increase in nornicotine concentrations in both undamaged top leaves of herbivore damaged plants and herbivore damaged leaves exposed to one and three larvae. The nicotine concentration increased in damaged compared to undamaged middle leaves. The nornicotine concentration was lower in damaged leaves of plants exposed to six compared to three larvae, suggesting that N. africana rather invests in new growth as opposed to protecting older leaves under severe attack. The results indicate that the nornicotine-rich N. africana will be unattractive to herbivores and more so when damaged, but that potential pollinators will be unaffected because the nectar remains alkaloid-free even after herbivory.     

光处理对白头翁愈伤组织生长及代谢产物合成的影响

以白头翁试管苗叶片为外植体,于MS＋TDZ0.2 mg/L＋2,4-D 0.2 mg/L上进行愈伤组织诱导和增殖,进行不同光质照射,观察光处理下白头翁愈伤组织的生长和代谢产物合成情况.结果表明：黄光能提高愈伤组织的诱导率,产生的愈伤组织质地最好;光质对愈伤组织的鲜重增殖倍数依次为：黄（16.69）〉 红（11.41）〉 绿（8.97）〉 白（6.74）〉蓝（5.97）,延长光照时间会促进细胞生长;与白光相比,黄光和红光能显著促进干物质的积累,而蓝光明显抑制,绿光与白光两处理之间差异不明显.自然光照下,白头翁叶片中代谢产物极少,而黄、蓝和绿光均诱导合成了包括皂苷在内的数种代谢产物,而红光处理同白光一样只诱导出了一种代谢产物.由此可见,光处理可以调控白头翁愈伤组织的生长状态及皂苷等次生代谢物的积累.     

Not so simple after all: searching for ecological advantages of compound leaves

Leaves come in many sizes and shapes, and the relationships between leaf traits and the environments they occur in are better understood every day. However we still know very little about the ecological consequences of plants having either compound or simple leaves. We attempted to address this knowledge gap by comparing chemical and physical characteristics (leaf area, length:width ratio, water content, leaf mass per area, ‘toughness’ and C:N ratio), as well as rates of herbivory between compound and simple leaves across 34 species in adjacent rainforest, open woodland and wet sclerophyll (tall open forest) vegetation in northeastern Australia. We found C:N ratio to be lower in simple leaves, but this was the only leaf trait that differed significantly between leaf types and did not stand up under phylogenetic analysis. Overall, we found no differences in herbivory between simple and compound leaves. While it remains unclear what the advantages of having one leaf type over another might be, the differences do not seem to lie in construction, or in vulnerability to herbivores, at least in the Australian Wet Tropics.     

Red (anthocyanic) leaf margins do not correspond to increased phenolic content in New Zealand Veronica spp.

Background and Aims

Red or purple coloration of leaf margins is common in angiosperms, and is found in approx. 25 % of New Zealand Veronica species. However, the functional significance of margin coloration is unknown. We hypothesized that anthocyanins in leaf margins correspond with increased phenolic content in leaf margins and/or the leaf entire, signalling low palatability or leaf quality to edge-feeding insects.

Methods

Five species of Veronica with red leaf margins, and six species without, were examined in a common garden. Phenolic content in leaf margins and interior lamina regions of juvenile and fully expanded leaves was quantified using the Folin–Ciocalteu assay. Proportions of leaf margins eaten and average lengths of continuous bites were used as a proxy for palatability.

Key Results

Phenolic content was consistently higher in leaf margins compared with leaf interiors in all species; however, neither leaf margins nor more interior tissues differed significantly in phenolic content with respects to margin colour. Mean phenolic content was inversely correlated with the mean length of continuous bites, suggesting effective deterrence of grazing. However, there was no difference in herbivore consumption of red and green margins, and the plant species with the longest continuous grazing patterns were both red-margined.

Conclusions

Red margin coloration was not an accurate indicator of total phenolic content in leaf margins or interior lamina tissue in New Zealand Veronica. Red coloration was also ineffective in deterring herbivory on the leaf margin, though studies controlling for variations in leaf structure and biochemistry (e.g. intra-specific studies) are needed before more precise conclusions can be drawn. It is also recommended that future studies focus on the relationship between anthocyanin and specific defence compounds (rather than general phenolic pools), and evaluate possible alternative functions of red margins in leaves (e.g. antioxidants, osmotic adjustment).     

Spatial patterning of pigmentation in evergreen leaves in response to freezing stress

Evergreen leaves of temperate climate plants are often subject to frosts. Changes in carbon gain patterns arise from freezing‐related tissue damage, and from interactions between light and temperature stress. We examined relationships between spatial patterns in freezing and concentrations of chlorophyll. Spatial patterns in pigmentation in leaves that had or had not been exposed to naturally occurring frosts were determined by conventional extraction techniques combined with high‐resolution hyperspectral imaging of reflectance from intact leaves. Predictive indices were developed to relate reflectance to chlorophyll content and chlorophyll a/b ratios within intact leaves. Leaves exposed to frosts had lower chlorophyll contents and more variable a/b ratios than protected leaves. In frost‐affected leaves, chlorophyll content was highest near leaf centres and decreased toward leaf tips and margins. Decline in chlorophyll content was associated with shifts in chlorophyll a/b ratios and increases in red pigmentation due to anthocyanin, with effects being greater on leaf sides exposed directly to the sun. These altered pigmentation patterns were consistent with patterns in freezing. The present results illustrate the fine scale of spatial variation in leaf response to freezing, and raise important questions about impacts of freezing on photosynthetic function in over‐wintering evergreens.     

Colour discrimination in dim light by the larvae of the African catfish <Emphasis Type="Italic">Clarias gariepinus</Emphasis>

Many demersal fish species undergo vertical shifts in habitats during ontogeny especially after larval metamorphosis. The visual spectral sensitivity shifts with the habitat, indicating a change in colour vision. Colour vision depends on sufficient ambient light and becomes ineffective at a particular low light intensity. It is not known how fishes see colour in dim light. By means of a behavioural experiment on larval African catfish Clarias gariepinus in the laboratory, we determined colour vision and colour discrimination in dim light. Light-adapted larvae were subjected to classical conditioning to associate a reward feed with a green or a red stimulus placed among 7 shades of grey. The larvae learned this visual task after 70 and 90 trials. A different batch of larvae were trained to discriminate between green and red and then tested for the ability to discriminate between these colours, as the light intensity was reduced. The larvae learned this visual task after 110 trials in bright light and were able to discriminate colours, as light was dimmed until 0.01 lx, the minimal illuminance measurable in this study, and similar to starlight. The retinae of the larvae were found to be light adapted at 0.01 lx; thus indicating cone-based colour vision at this illuminance. For comparison, three human subjects were tested under similar conditions and showed a colour vision threshold at between 1.5 and 0.1 lx. For the larvae of C. gariepinus, the ability of colour discrimination in dim light is probably due to its retinal tapetum, which could increase the sensitivity of cones.     

常绿阔叶林中栲树展叶期叶片的虫食格局

叶片的虫食主要发生在展叶期。虽然展叶期短暂, 它却可能是了解植食性昆虫和植物之间相互关系的关键。为了解栲树(Castanopsis fargesii)在展叶期叶片的虫食格局和展叶方式对叶片虫食的影响, 研究了栲树展叶期内的虫食动态变化, 结果表明: 栲树展叶的两个阶段(折叠期和打开期), 虫食叶片的格局存在较大的差异, 打开阶段的日虫食频度和日虫食率显著高于折叠阶段(F_{1, 32}=8.97, p=0.005 4; F_{1, 32}=12.38, p=0.001 4), 展叶期最终叶片虫食频度为50.72%, 叶片虫食率为8.25%。折叠期叶片主要受到低强度的虫食, 打开期叶片虫食则以较大强度的虫食为主。展叶期叶片的虫食主要发生在夜间, 夜间虫食率显著高于日间虫食率(t=2.51, p=0.017), 变化趋势与日虫食率一致。栲树叶片在展叶的两个阶段可能采用了不同的防御对策。     

Functional role of red (retro)-carotenoids as passive light filters in the leaves of Buxus sempervirens L.: increased protection of photosynthetic tissues?

Red (retro)-carotenoids accumulate in chloroplasts of Buxus sempervirens leaves during the process of winter leaf acclimation. As a result of their irregular presence, different leaf colour phenotypes can be found simultaneously in the same location. Five different colour phenotypes (green, brown, red, orange, and yellow), with a distinct pattern of pigment distribution and concentration, have been characterized. Leaf reddening due to the presence of anthocyanins or carotenoids, is a process frequently observed in plant species under photoinhibitory situations. Two main hypotheses have been proposed to explain the function of such colour change: antioxidative protection exerted by red-coloured molecules, and green light filtering. The potential photoprotective role of red (retro-) carotenoids as light filters was tested in Buxus sempervirens leaves. In shade leaves of this species the upper (adaxial) mesophyll of the lamina was replaced by the equivalent upper part of a different colour phenotype. These hybrid leaves were exposed to a photoinhibitory treatment in order to compare the photoprotective effect exerted by adaxial parts of phenotypes with a different proportion of red (retro)-carotenoids in the lower mesophyll of a shade leaf. The results indicated that the presence of red (retro)-carotenoids in the upper mesophyll did not increase photoprotection of the lower mesophyll when compared with chlorophyll, and the best protection was achieved by an upper green layer. This was due to the fact that the extent of photoinhibition was proportional to the amount of red light transmitted by the upper mesophyll and/or to the chlorophyll pool located above. These results do not exclude a protective function of carotenoids in the upper leaf layer, but imply that, at least under the conditions of this experiment, the accumulation of red pigments in the outer leaf layers does not increase photoprotection in the lower mesophyll.     

有色膜遮光对烤烟生长和光合特性及其初烤品质的影响

以自然光为对照,采用红色、白色、蓝色、黄色4种有色薄膜于2010~2011年从团棵期开始对大田烤烟进行遮光处理,研究不同光质对烤烟生长、光合特性及初烤品质指标的影响。结果显示:(1)红膜处理最大叶长宽比最小、叶面积最大,黄膜处理则相反。(2)红、蓝膜处理烟叶净光合速率、气孔导度、蒸腾速率明显高于自然光处理,白、黄膜处理略高于对照或与对照持平,且遮膜处理前期红膜高于蓝膜处理,后期蓝膜高于红膜处理。(3)红、蓝膜处理有利于提高倒5叶SPAD值,黄膜处理则相反。(4)红膜处理显著降低了中部叶蛋白质、总氮含量和氮碱比,提高了施木克值,并显著提高了上部叶可溶性糖含量和氮碱比,降低了施木克值;蓝膜处理显著提高了中部叶烟碱和多酚含量,降低了可溶性糖含量、施木克值及氮碱比,并显著提高了上部叶蛋白质、总氮、烟碱和多酚含量,降低了施木克值,提高了氮碱比;黄膜处理显著降低了中上部叶蛋白质、总氮、烟碱和多酚含量,提高了上部叶施木克值、降低了氮碱比。研究表明,红、蓝膜处理更利于烟叶发育和光合特性的提高,初烤烟叶化学成分更协调,利于优质烟叶的形成。     

Morphology and Photosynthetic Efficiency of Tobacco Leaves That Received End-of-Day Red and Far Red Light during Development

Shaded leaves in plant canopies receive a higher proportion of far red relative to red light than is received by unshaded leaves. Brief end-of-day irradiations with red or far red light, acting through the phytochrome system, reversibly control morphological development of tobacco plants. Leaves that received far red light for 5 minutes at the end of each day during development were longer and narrower than those that received end-of-day red light. The far red treated leaves weighed less, had fewer stomata, and had less chlorophyll per unit area of leaf. Net CO₂ assimilation rates did not differ significantly between red- and far red-treated leaves on an area basis; however, the far red-treated leaves assimilated significantly more CO₂ on a leaf weight basis.     

Herbivory pattern and induced responses to simulated herbivory inQuercus mongolica var.grosseserrata

Seasonal changes in leaf traits and the herbivory pattern ofQuercus mongolica var.grosseserrata were studied, and simulated herbivory experiments were carried out in order to evaluate leaf trait responses. Leaves ofQ. mongolica emerged simultaneously in spring and most were retained until autumn. Nitrogen concentration was highest when leaves first emerged and decreased rapidly with leaf age. Leaf mass per area (LMA) increased with leaf age. Herbivore attack was concentrated in the first 20 days after bud-break, which corresponded to the high nutritional value of the leaves for herbivores at this time. Simulated herbivory experiments indicated that LMA increased with artificial leaf damage, suggesting an increase in leaf toughness, and that nitrogen concentration decreased later in the season in comparison with intact leaves. As a result, herbivore attack following artificial leaf damage decreased with increasing initial leaf damage. However, leaf longevity was not affected by initial leaf damage. These responses were considered to be a strategy to disperse herbivory damage among leaves.     

Photomorphogenesis, Photosynthesis, and Early Growth of Primary Leaves of Phaseolus vulgaris

The response of leaf tissue to white, blue, red, and far-redlight has been examined. Leaves on plants grown in darknessshow increased cell number, cell volume, and area when exposedto long periods (up to 48 h) of low-intensity red, blue, orfar-red radiation. This is believed to be a photomorphogenicresponse which does not involve photosynthesis. Leaves fromplants exposed to white light during germination do not usuallyrespond to red, blue, or far-red light. An exception to thiswas found for leaf discs which showed a larger increase in areathan the dark controls following exposure to far-red light for24 h. Leaf tissue from light-grown plants responds to high-intensitywhite light, probably through photosynthesis. Discs cut fromdark-grown plants and cultured in white light grow equally wellin air and CO₂-free conditions. Application of the photosyntheticinhibitor DCMU reduces growth and chlorophyll formation, however. It is concluded that light, perhaps acting through the phytochromemechanism, has initially a number of morphogenic effects includinginitiation of development of the photosynthetic apparatus. Theresponses to photomorphogenically active radiation do not persistand light effects through photosynthesis are rapidly initiatedand dominate the later stages of leaf growth.     

Do small leaves expand faster than large leaves,and do shorter expansion times reduce herbivore damage?

Leaves are most vulnerable to herbivory during expansion. We hypothesised that one factor favouring small leaves could be that smaller-leaved species have shorter expansion times and are therefore exposed to high levels of herbivory for a shorter period than large leaves. In order to test this hypothesis, leaf expansion time and leaf area loss were measured for 51 species from Sydney, Australia. Strong positive correlations were found between leaf length and area and leaf expansion time, confirming that small leaves do expand in a shorter time than large leaves. The amount of leaf area lost was highly variable (from 0.5 to 90% of total leaf area), but was significantly related to both leaf expansion time and log leaf area. The amount of leaf area lost was not significantly correlated with specific leaf area nor with the presence of distasteful substances in the leaves, but was lower on species with hairy expanding leaves.     

Differential induced response to generalist and specialist herbivores by Lindera benzoin (Lauraceae) in sun and shade

Theoretically, induced defenses should be prevalent within low resource environments like the forest understory where constitutive defenses would be costly. Also, the induced response should be stronger when the herbivore is a generalist rather than a specialist, which often have mechanisms to avoid or overcome plant defenses. These ideas have been previously tested for herbaceous species, and we examined these predictions in Lindera benzoin (spicebush), a common woody shrub of the eastern deciduous forest. Lindera benzoin plants in contrasting light environments served as control plants or were subjected to one of four treatments: application of jasmonic acid, clipping, herbivory by the specialist Epimecis hortaria (tulip tree beauty) and herbivory by the generalist Spodoptera exigua (beet armyworm). Following treatment, we assessed induced responses by measuring leaf chemistry (C/N ratio, protein content, and peroxidase activity), and by using insect bioassays with E. hortaria larvae. We found no difference in peroxidase activity between light environments in controls, plants treated with clipping or jasmonic acid. In plants subject to insect herbivory, peroxidase activity was greater in shade plants than in sun plants. The magnitude of this increase in the shade varied between the herbivores, with a 32 fold increase in plants exposed to the generalist S. exigua and a 9 fold increase in plants exposed to the specialist E. hortaria . Leaves from shade plants had more protein and lower C/N ratios than leaves from sun plants, regardless of induction treatment. In control plants, E. hortaria larvae consumed more leaf biomass and achieved greater final weights in the sun than in the shade, but these differences disappeared with induction treatments were applied. These results are among the first to show rapid induction in a woody plant, and different levels of induction with light environments and with specialist versus generalist herbivores.     

Herbivory limits the yellow water lily in an overgrown lake and in flowing water

Aquatic macrophytes with floating leaves are often key ecological species that affect entire aquatic ecosystems. Here we describe an investigation of the importance of insect herbivory for population growth and leaf senescence in the yellow water lily (Nuphar lutea). In order to gain a general picture of the importance of herbivory under different conditions, we experimentally manipulated herbivory in a large lily population in natural still water and observed the natural development of 32 smaller populations in flowing water. Herbivory drastically increased leaf senescence, reducing leaf density. In the still water, over one summer, leaf density increased by a factor of 1.23 in the presence of water lily leaf beetles and 1.61 when herbivory was eliminated. In flowing water, population growth was restricted mainly by leaf crowdedness, which limited large dense populations. Herbivory by water lily leaf beetles also had a limiting effect on yellow water lily, again mainly in large dense populations. Small populations supported a lower density of beetles. Previous studies have not addressed population-level responses of vascular plants with floating leaves. Our results suggest that herbivory can result in greater light penetration into the water and reduce “enemy-free space” for aquatic species that find such space in water lily stands. We suggest that the water lily leaf beetle should be considered an “ecological engineer.”