Injured and uninjured leaf photosynthetic responses after mechanical injury on <Emphasis Type="Italic">Nerium oleander</Emphasis> leaves,and <Emphasis Type="Italic">Danaus plexippus</Emphasis> herbivory on <Emphasis Type="Italic">Asclepias curassavica</Emphasis> leaves

Insect herbivory has variable effects on plant physiology; so greater understanding is needed about how injury alters photosynthesis on individual injured and uninjured leaves. Gas exchange and light-adapted leaf chlorophyll fluorescence measurements were collected from uninjured and mechanical partial leaf defoliation in two experiments with Nerium oleander (Apocynaceae) leaves, and one experiment with Danaus plexippus herbivory on Asclepias curassavica (Asclepiadaceae) leaves. Gas exchange impairment (lower photosynthetic rate (P _n ), stomatal conductance (g _s)) indicates water stress in a leaf, suggests stomatal limitations causing injury P _n impairment. The same pattern of gas exchange impairment also occurred on uninjured leaves opposite from injured leaves in both N. oleander experiments. This is an interesting result because photosynthetic impairment is rarely reported on injured leaves near injured leaves. No photosynthetic changes occurred in uninjured A. curassavica leaves opposite from D. plexippus-fed leaves. Partially defoliated leaves that had P _n and g _s reductions lacked any significant changes in intercellular leaf [CO₂], C _i. These results neither support, nor are sufficient to reject, stomatal limitations to photosynthesis. Manually imposed midrib vein severance in N. oleander experiment #1 significantly increased leaf C _i, indicating mesophyll limitations to photosynthesis. Maximal light-adapted leaf photochemical efficiency () and also non-photochemical quenching (q _N) were reduced by mechanical or insect herbivory to both study species, suggesting leaf trouble handling excess light energy not used for photochemistry. Midrib injury on N. oleander leaves and D. plexippus herbivory on A. curassavica leaves also reduced effective quantum yield (ΦPSII) and photochemical quenching (q _P); so reduced plastoquinone pools could lead to additional PSII reaction center closure.     

Developmental plasticity in birch leaves: defoliation causes a shift from glandular to nonglandular trichomes

The structures on leaf surfaces, e.g. trichomes, can act as effective antiherbivory mechanisms as chemical repellents. Structural defences usually represent constitutive resistance, but there are also a few cases of inducible morphological defences. We tested whether defoliation may induce changes in trichome production in white birch (Betula pubescens). The studied birches were either 0, 50 or 100% defoliated during the previous or current summer, and we measured the alterations in the production of glandular vs. nonglandular leaf trichomes, developmental instability (fluctuating asymmetry, FA) and leaf and shoot growth. We detected a clear shift from glandular to nonglandular leaf trichomes following previous‐year defoliation but not after current‐year defoliation. Furthermore, the density of nonglandular trichomes around the mid‐vein of leaves increased following previous‐year defoliation but decreased after current‐year defoliation. While leaf and shoot growth showed a distinct decrease in response to defoliation, FA turned out to be less sensitive. Consequently, previous‐year defoliation can induce the production of nonglandular trichomes in birch leaves. Because this response was accompanied by a reduction in glandular trichomes, the present results may suggest a trade‐off between the different trichome types of birch leaves.     

Effects of the experimental blockage of the major veins on hydraulics and gas exchange of Prunus laurocerasus L. leaves

The impact of leaf vein blockage on leaf hydraulic conductance (K(L)), gas exchange (g(L)) and water potential (Psi(L)) was studied in Prunus laurocerasus L., a broad-leaved evergreen. For this purpose, leaves were measured for the three variables above, either with an intact leaf blade (controls) or with the midrib cut a third of the way up (cut a), or with the midrib cut at three different points and the first-order veins cut through near their insertion to the midrib (cut b), or with the midrib cut at 2 mm from the leaf base (cut c). All the cut surfaces were sealed with cyanoacrylate. A serial decrease of K(L) was recorded from cut a to cut c with respect to that measured for the controls, i.e. a K(L) loss of about 37% (cut a), 57% (cut b) and 87% (cut c). A positive linear relationship appeared to exist between g(L) and K(L) with a high correlation coefficient (r(2)=0.99) and a high statistical significance (P <0.01). Even under a severe drop in K(L) (as that induced by cut c), leaf water potential remained approximately constant and not statistically different from Psi(L) measured for the controls. In fact, Psi(L) ranged between -0.83 and -0.98 MPa, i.e. within the cavitation threshold of leaves in terms of the critical Psi(L) inducing a significant production of ultrasound acoustic emissions which was -0.94+/-0.09 MPa. The conclusion was that stomata were very sensitive to changes in K(L) and that stomatal closure led to the homeostatic maintenance of Psi(L) and cavitation avoidance.     

浙江天童常绿阔叶林石栎叶片昆虫取食状类型和取食强度研究

在浙江天童常绿阔叶林中随机选择10株石栎（Lithocarpus glaber）大树,每株选择冠层向阳枝条2枝,取其所有成熟叶片（2332片）对昆虫取食状类型和取食强度进行了分析。结果表明,石栎叶片昆虫取食状类型多样,计有15种,但出现频度有很大不同;顶食状和缘食状的频度分别在30％以上;切叶状、连续小孔状和大孔状的频度在10％～30％：小孔状、阴面食状、叶中脉、泡状的频度在6．4％～1．5％;其余6种类型的频度〈1％。对叶片的危害程度取决于取食面积和频度,顶食状等3种类型频度高、取食面积大,对叶片的危害重;而阴面食状等11种类型取食面积较小,频度低,对叶片的危害较小;连续小孔状尽管取食频度高,但取食面积小,对叶片的危害也小。连续小孔状和切叶状之间存在显著正相关,但连续小孔食状与阴面食状之间,切叶状、连续小孔食状与小孔食状之间则存显著负相关,其它取食状之间均没有显著相关性,反映出不同昆虫类型的取食生态位关系。石栎叶片昆虫取食率为13．6％,介于温带森林和热带森林之间;但石栎叶片的昆虫取食频率64％,小于辽东栎,可能是由于防御机制上的差异所至;此外,石栎叶片昆虫取食强度还受到植株本身因素和小生境等因素的影响。     

Decline of leaf hydraulic conductance with dehydration: relationship to leaf size and venation architecture

Across plant species, leaves vary enormously in their size and their venation architecture, of which one major function is to replace water lost to transpiration. The leaf hydraulic conductance (K(leaf)) represents the capacity of the transport system to deliver water, allowing stomata to remain open for photosynthesis. Previous studies showed that K(leaf) relates to vein density (vein length per area). Additionally, venation architecture determines the sensitivity of K(leaf) to damage; severing the midrib caused K(leaf) and gas exchange to decline, with lesser impacts in leaves with higher major vein density that provided more numerous water flow pathways around the damaged vein. Because xylem embolism during dehydration also reduces K(leaf), we hypothesized that higher major vein density would also reduce hydraulic vulnerability. Smaller leaves, which generally have higher major vein density, would thus have lower hydraulic vulnerability. Tests using simulations with a spatially explicit model confirmed that smaller leaves with higher major vein density were more tolerant of major vein embolism. Additionally, for 10 species ranging strongly in drought tolerance, hydraulic vulnerability, determined as the leaf water potential at 50% and 80% loss of K(leaf), was lower with greater major vein density and smaller leaf size (|r| = 0.85-0.90; P < 0.01). These relationships were independent of other aspects of physiological and morphological drought tolerance. These findings point to a new functional role of venation architecture and small leaf size in drought tolerance, potentially contributing to well-known biogeographic trends in leaf size.     

Novel fungal disease in complex leaf‐cutting ant societies

Abstract 1. The leaf‐cutting ants practise an advanced system of mycophagy where they grow a fungus as a food source. As a consequence of parasite threats to their crops, they have evolved a system of morphological, behavioural, and chemical defences, particularly against fungal pathogens (mycopathogens). 2. Specific fungal diseases of the leaf‐cutting ants themselves have not been described, possibly because broad spectrum anti‐fungal defences against mycopathogens have reduced their susceptibility to entomopathogens. 3. Using morphological and molecular tools, the present study documents three rare infection events of Acromyrmex and Atta leaf‐cutting ants by Ophiocordyceps fungi, agenus of entomopathogens that is normally highly specific in its host choice. 4. As leaf‐cutting ants have been intensively studied, the absence of prior records of Ophiocordyceps suggests that these infections may be a novel event and that switching from one host to another is possible. To test the likelihood of this hypothesis, host switching was experimentally induced, and successfully achieved, among five distinct genera of ants, one of which was in a different sub‐family than the leaf‐cutter ants. 5. Given the substantial differences among the five host ants, the ability of Ophiocordyceps to shift between such distant hosts is remarkable; the results are discussed in the context of ant ecological immunology and fungal invasion strategies.     

A comparison of artificial defoliation techniques using canola (<Emphasis Type="Italic">Brassica napus</Emphasis>)

We conducted two experiments that investigated how the method and location of artificial defoliation influenced growth, reproduction, and allocation in canola, Brassica napus. In one experiment, 0%, 25%, or 50% of leaf area was removed by cutting circular holes at three possible locations: concentrated at either the base of leaves or at their tips, or dispersed throughout leaf blades. Plants fully compensated for such damage; reproduction and allocation were unaffected by either defoliation intensity or wound location. In a second experiment, we again initiated three intensities of defoliation: non-damaged plants served as controls, while others had 25% or 50% of their leaf areas removed. The method of removal in the second experiment consisted of cutting either multiple, similar-sized, circular holes or single, contiguous patches of a leaf blade. At the highest defoliation intensity reproductive output and allocation were significantly less in plants treated with the former method than the latter, even though an equivalent initial amount of leaf area was removed in both treatments. We conclude that simulated herbivory studies must account for not only how much of the plant is damaged, but also the pattern of leaf damage itself, since both factors contribute to a plant’s physiological and ecological responses to grazing.     

Photosynthetic responses of potato to Colorado potato beetle injury and differences in injury between adult males and females

Establishing rates of injury to plants and the physiological impact of this injury provides essential data in the development of economic injury levels, but variation of sex effects is not often considered. Here, we examined injury by the Colorado potato beetle, Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), larvae and adult males and females on potato, Solanum tuberosum L. (Solanaceae). Specifically, we looked for adult sex differences between males and females in injury rates (= leaf consumption rates), and examined the impact of all types of injury (larval, adult male, and adult female) on gas exchange parameters of remaining potato leaf tissue. Experiments were conducted in the field and in growth chambers on Frito‐Lay proprietary and Pike chipping‐potato varieties at pre‐blooming and blooming stages. We found no change in photosynthetic rates on remaining (uninjured) leaf tissue infested with male, female, or fourth‐stage larva of Colorado potato beetle. However, when the midrib was cut in trials with male beetles, the remaining tissue above the injury exhibited photosynthetic rate reductions as a result of stomatal limitations. These findings are consistent with the pattern that we and other researchers have observed with gross tissue removal by various insects on other plant species. Adult females consumed more tissue than males, and temperature was positively correlated with feeding rates for both sexes. Sex‐related differences in feeding rate are most important to studies quantifying consumption rates for economically important species because of its potential impact on resulting economic injury level calculations.     

Grain Amaranths Are Defoliation Tolerant Crop Species Capable of Utilizing Stem and Root Carbohydrate Reserves to Sustain Vegetative and Reproductive Growth after Leaf Loss

Tolerance to defoliation can be defined as the degree to which productivity is affected by photosynthetic area reduction. This trait was studied in grain amaranth (Amaranthus cruentus and A. hypochondriacus), which are considered to be a highly defoliation-tolerant species. The physiological and biochemical responses to increasing levels of mechanical leaf removal up to total defoliation were quantified. Tolerance appeared to be dependent on various factors: ( i) amount of lost tissue; (ii) mechanics of leaf tissue removal; (iii) environment, and (iv) species tested. Thus, grain amaranth was found to be a highly tolerant species under green-house conditions when leaf tissue loss was performed by gradual perforation. However, tolerance was compromised under similar conditions when defoliation was done by gradual cutting of the leaf. Also tolerance in completely defoliated plants tended to decrease under field conditions, where differences between A. cruentus and A. hypochondriacus were observed. All non-structural carbohydrate (NSC) levels were reduced in stems and roots of totally defoliated amaranths one day after treatment. Such depletion probably provided the carbon (C) resources needed to sustain the early recovery process in the absence of photosynthetic capacity. This was corroborated by shading of intact plants, which produced the same rapid and drastic reduction of NSC levels in these tissues. These results emphasize the role of stored NSCs, particularly starch, in buffering the impact of severe defoliation in amaranth. The fall in sucrose synthase and cell wall invertase activity observed in stems and roots soon after defoliation was consistent with their predicted shift from sink to source tissues. It is concluded that mobilization of C stores in stems and roots, is a physiologically important trait underlying tolerance to defoliation in grain amaranth.     

Effect of Excision on Leaf Water Potential

Improvements of thermocouple hygrometric techniques for in situleaf water potential measurement in the field now allows forcontinuous monitoring of water potential in response to an externalperturbation, such as leaf excision. Using Citrus jambhiri plants,measured leaf water potentials of completely excised leaf portionsimmediately increased when the petiole was excised or incisionswere made either transverse or parallel to the midrib. Incisionsparallel to the midrib were on the side nearest the hygrometeror opposite it if preceded by a petiole excision. Midrib incisionswere 100–150 mm long with the nearest cut edge being 20–50mm from the hygrometer cavity. All excisions were such thatleaf tissue was removed from the leaf with water potential onone of the leaf portions being measured continuously prior toand after excising. The peak increase in measured water potentialof the excised leaf portions ranged between 20 kPa and 80 kPabut averaged 50 kPa. In uncovered leaves, particularly underfield conditions with the associated high evaporative demand,measured leaf water potential declined rapidly after the initialincrease. The increase in measured water potential immediatelyfollowing various types of excision was confirmed for dark andlight conditions (laboratory and field respectively) using bothpsychrometric and dewpoint modes and occurred for secondaryexcisions, but to a lesser extent. Discovery of this phenomenonimplies that water potential measured on detached leaves maynot always represent accurately in situ leaf water potential. Key words: Leaf water potential, Thermocouple hygrometers, Leaf excision effects     

一号冰川地区四种藓类植物的解剖学研究

运用石蜡切片技术和扫描电镜方法,对一号冰川地区生长的4种藓类植物茎、叶的内部结构及叶表皮角质层褶皱、疣和中肋等进行观察,结果表明：红扭口藓(Barbula asperifolia Mitt.)茎横切面呈多棱形;叶背、腹面角质层厚,粗疣不分叉,但顶端朝细胞凹陷处倾斜;叶背面细胞壁凹陷深,象张开的气孔,粗疣也藏在其中,中肋突出明显。丛叶扭藓(Tortella humilis (Hedw.) Jenn.)茎横切面呈椭圆形;叶背、腹面中上部均密被鹿角状粗疣,这些分叉的粗疣,从凹陷的细胞壁处成束突起,顶端向下弯曲成钩状,在叶的下部疣状突起则逐渐减少至无,中肋较宽。异叶提灯藓(Mnium heterophyllum (Hook.)Schwagr.)茎横切面呈五棱形;叶细胞一层,呈不规则多边形,细胞壁凹陷使叶片呈网状;中肋红色。大灰藓(Hypnum plumaeforme Wils.)茎的横切面呈圆形;叶背、腹面均密被疣状突起,且差异不大,粗疣顶部均倒向孔口呈遮盖状;中肋短而弱。     

Hydraulic analysis of water flow through leaves of sugar maple and red oak

Leaves constitute a substantial fraction of the total resistance to water flow through plants. A key question is how hydraulic resistance within the leaf is distributed among petiole, major veins, minor veins, and the pathways downstream of the veins. We partitioned the leaf hydraulic resistance (R(leaf)) for sugar maple (Acer saccharum) and red oak (Quercus rubra) by measuring the resistance to water flow through leaves before and after cutting specific vein orders. Simulations using an electronic circuit analog with resistors arranged in a hierarchical reticulate network justified the partitioning of total R(leaf) into component additive resistances. On average 64% and 74% of the R(leaf) was situated within the leaf xylem for sugar maple and red oak, respectively. Substantial resistance-32% and 49%- was in the minor venation, 18% and 21% in the major venation, and 14% and 4% in the petiole. The large number of parallel paths (i.e. a large transfer surface) for water leaving the minor veins through the bundle sheath and out of the leaf resulted in the pathways outside the venation comprising only 36% and 26% of R(leaf). Changing leaf temperature during measurement of R(leaf) for intact leaves resulted in a temperature response beyond that expected from changes in viscosity. The extra response was not found for leaves with veins cut, indicating that water crosses cell membranes after it leaves the xylem. The large proportion of resistance in the venation can explain why stomata respond to leaf xylem damage and cavitation. The hydraulic importance of the leaf vein system suggests that the diversity of vein system architectures observed in angiosperms may reflect variation in whole-leaf hydraulic capacity.     

CHARACTERIZATION OF DEVELOPMENT IN MAIZE THROUGH THE USE OF MUTANTS. II. THE ABNORMAL GROWTH CONDITIONED BY THE KNOTTED MUTANT

The maize mutant Knotted (Kn) is characterized by hollow, finger-like outgrowths (knots) occurring mainly in the leaf blade. Portions of the ligule are displaced from the normal position to more distal locations within the blade. Knots apparently result from continued meristematic activity of isolated patches of cells surrounded by maturing tissue. Small knots appear to be centers of cell division. Epidermal cells overlying a small knot have been observed to undergo periclinal divisions. In addition to cell division, a reorientation of the axis of cell elongation is associated with knot formation. The pattern of knot distribution varies at different levels on the plant axis and within a leaf blade. From leaf 4 to leaf 10 or 11 the number of knots per leaf increases progressively, then declines in leaves initiated later. Knots always occur in association with lateral veins. The greatest number per vein occurs on the 3rd or 4th vein from the midrib. One plant developing from an X-rayed heterozygous seed possessed a sector of normal tissue bisecting the plant in a vertical plane passing through the midrib of each leaf except the top two. The normal sector was knot-free and had the ligule restored to the normal position. These observations suggest that cells with the characteristics of those from intercalary meristems occur throughout the blade in Knotted plants.     

Pattern of defoliation and its effect on photosynthesis and growth of Goldenrod

1. Leaf area was removed from Solidago altissima in either a dispersed pattern (half of every leaf removed) or a concentrated pattern (every other leaf removed) and effects on leaf gas exchange, vegetative growth and flowering were examined relative to undefoliated controls. Gas exchange was measured for leaves remaining after defoliation and for regrowth leaves that developed post-damage (at 7, 16 and 26 days post-defoliation).
2. Area-based photosynthetic rates of leaves remaining after defoliation were not affected by either dispersed or concentrated damage, but damage of both types enhanced area-based photosynthesis of regrowth leaves at 16 days post-defoliation and to a lesser extent at 26 days post-defoliation.
3. Dispersed damage, but not concentrated damage, stimulated mass-based photosynthesis of undamaged leaves remaining after defoliation. Undamaged leaves remaining after defoliation and regrowth leaves on damaged plants had higher specific leaf area (leaf area/leaf mass) than comparable leaves on control plants. Mass-based photosynthesis was more strongly elevated by defoliation than area-based photosynthesis because of this increase in specific leaf area.
4. Plants with dispersed damage recovered more quickly from defoliation; they had higher relative growth rates in the first week post-defoliation than plants with concentrated damage. Both types of defoliation caused similar reductions in flower production.
5. These results add to accumulating evidence that dispersed damage is generally less detrimental to plants than concentrated damage and suggest that physiological changes in leaves may be part of the reason.     

EVOLUTION OF DIFFERENT GALL TYPES IN WILLOW-FEEDING SAWFLIES (HYMENOPTERA: TENTHREDINIDAE)

The sawflies that feed on the plant family Salicaceae can be divided into eight informal groups based on larval feeding habit or gall type: (1) species with free-living larvae; (2) leaf folders; (3) leaf blade gallers; (4) apical leaf gallers; (5) basal leaf gallers; (6) midrib and petiole gallers; (7) stem gallers; and (8) bud gallers. It has been proposed that the galling habit evolved from free-living larvae via leaf folders, and that the different gall types evolved gradually in the sequence mentioned above. Thus, the galling site would have “wandered” from the leaf margin toward the stem as a result of gradual changes in oviposition site preference. Allozyme data from eight informative loci were used to reconstruct the phylogeny of 18 representative sawfly species. The results suggest that indeed leaf folders seem to be a basal group; leaf blade gallers evolved independently of the other true gallers; apical and basal leaf gallers are not the ancestors of petiole and bud gallers, but they may share a common galling ancestor; bud gallers evolved from midrib/petiole gallers; and stem gallers are polyphyletic. The cause for the observed wandering of the galling site could be intraspecific competition due to a possible “nutrient shading effect” of galls situated closer to the host plant's main vascular system.     

Photosynthetic responses of wheat, Triticum aestivum L., to defoliation patterns on individual leaves

The impact of defoliation by fall armyworm, Spodoptera frugiperda (J. E. Smith), on the photosynthetic rates of injured, individual wheat, Triticum aestivum L., leaves and the impact of different spatial patterns of artificial insect defoliation on photosynthesis of remaining leaf tissue of injured, individual wheat leaves were evaluated in this study. Photosynthesis, stomatal conductance, transpiration, and chlorophyll a fluorescence were recorded in the flag-leaves of wheat plants 1 and 24 h after defoliation in 2003 and at 1 h, 24 h, 7 d, and 14 d after defoliation in 2004. Photosynthesis of injured leaves was not significantly affected by any defoliation treatment (i.e., control, natural, and artificial). Similarly, we did not observe interactions between defoliation treatments and time after defoliation. Stomatal conductance was significantly affected by time after defoliation and by the interaction between defoliation treatment and time after defoliation. However, in general, our results showed that wheat responded similarly to insect defoliation and artificial defoliation, which, therefore, may be used to simulate leaf mass consumption. Spatial defoliation patterns had a significant effect on photosynthetic parameters of injured leaves, but responses were dependent on plant developmental stages. The chlorophyll a fluorescence data revealed no significant effects from any defoliation pattern on the photochemical efficiency of the injured leaf. No significant interactions between defoliation patterns and time after defoliation were observed. Our findings reveal that the spatial pattern of defoliation in wheat affects photosynthetic and other gas exchange responses, which suggests that when simulating insect defoliation in wheat, researchers need to be cognizant of the defoliation pattern to adequately simulate insect defoliation.     

Defoliation and Regrowth in the Graminaceous Plant: The Role of Current Assimilate

Infra-red gas analysis and a quantitative radiocarbon tracertechnique were used to measure photosynthesis, and the export,distribution and utilization of current assimilate in the regrowthof leaf tissue and the growth of stem and root of partially-defoliateduniculm barley plants. After defoliation, which removed allleaf tissue above the ligule of leaf 3, the rate of photosynthesisof the remaining two older leaves fell to 90–95 per centof that of control leaves, but they exported more of their assimilatedcarbon to meristems elsewhere in the plant during the first48 h after the defoliation. The level of export from the twoolder leaves began to decline when new leaf tissue regrew fromthe shoot apex, and fell below that of the control leaves 4days after defoliation. The two older leaves supplied the assimilateused in the regrowth of new leaf tissue immediately after defoliation:previously they had exported most of their assimilate to root.There was no evidence that ‘reserves’ were mobilizedto meet the needs of regrowth at leaf meristems or, indeed,of the growth in stem and root; current photosynthesis suppliedsufficient assimilate to account for all observed growth. Ingeneral, the plants responded to defoliation with a rapid andmarked re-allocation of assimilate from root to leaf meristems,with the result that root growth was severely retarded but newleaf tissue grew at 70–100 per cent of the rate observedin control plants.     

干旱胁迫下紫穗槐叶片解剖特征的变化

研究了重度、中度、轻度干旱胁迫条件下2年生紫穗槐幼苗叶片的形态解剖学特征变化.结果表明:随着干旱胁迫程度的增加,叶片厚度、上下表皮的厚度逐渐减小,栅栏组织和海绵组织的厚度也减小,而栅栏组织与海绵组织的比值、主脉维管束相对直径、主脉突起度与中脉厚度则增大.气孔面积随干旱程度的增加呈下降趋势,但气孔分布的密度增加.在干旱逆...