Evolution of virulence when transmission occurs before disease

Most models of virulence evolution assume that transmission and virulence are constant during an infection. In many viral (HIV and influenza), bacterial (TB) and prion (BSE and CWD) systems, disease-induced mortality occurs long after the host becomes infectious. Therefore, we constructed a model with two infected classes that differ in transmission rate and virulence in order to understand how the evolutionarily stable strategy (ESS) depends on the relative difference in transmission and virulence between classes, on the transition rate between classes and on the recovery rate from the second class. We find that ESS virulence decreases when expressed early in the infection or when transmission occurs late in an infection. When virulence occurred relatively equally in each class and there was disease recovery, ESS virulence increased with increased transition rate. In contrast, ESS virulence first increased and then decreased with transition rate when there was little virulence early in the infection and a rapid recovery rate. This model predicts that ESS virulence is highly dependent on the timing of transmission and pathology after infection; thus, pathogen evolution may either increase or decrease virulence after emergence in a new host.     

Construction of a bio-test for infecting red clover plants with Sclerotinia trifoliorum

Sclerotinia trifoliorum causes clover cancer in red clover crops. Clover cancer is difficult to control and completely resistant red clover varieties are not available. Breeding for resistant red clover varieties is being slowed down because little is known about the diversity of European S. trifoliorum populations and because of the lack of bio-tests that are useable in breeding programs. The first objective of this research was to develop a reliable high-throughput bio-test, useable in breeding programs. The second objective was to optimise another bio-test, based on isolated leaves, for more precise studies. First, we optimised a method for ascospore production of S. trifoliorum. Once produced, the ascospores were used to evaluate the effects of climate conditions, ascospore concentration and plant age on the high-throughput bio-test. For the bio-test on isolated leaves, the effects of infection method, incubation conditions, incubation period, ascospore concentration, leaf growth stage and mechanical damage were evaluated. In the high-throughput bio-test, disease levels rose with increasing ascospore concentration up to 20,000 spores/ml. The plant age had a small, yet significant effect on the disease level. For the isolated leaf bio-test, the most effective and most repeatable infection method was spraying of an ascospore suspension. Disease levels continued to increase with rising concentrations and incubation time did not interact with plant susceptibility levels. The youngest completely opened leaf yielded the most repeatable results. Both bio-tests were shown to be correlated and could be valuable instruments for breeding programs and for studying plant-pathogen interactions.     

Why do some nectar foragers perch and others hover while probing flowers?

Diurnal hawkmoths, Hemaris fuciformis, and bumblebees, Bombus pasquorum, were observed foraging for nectar in flowers of Viscaria vulgaris. The hawkmoths hovered in front of the flowers, while the bees perched on them. The hawkmoths had a faster probing rate than the bees, and consequently also had higher gross and net rates of energy gain. A model is presented that shows that hovering only yields a higher net rate of energy gain (NREG) than perching when nectar volumes are high due to low competition for the resource. The difference in NREG of perchers and hoverers decreases with an increase of competition, and eventually perching yields the highest NREG. This is an effect of the higher cost of hovering. The results suggest that hovering can only evolve as a pure evolutionarily stable strategy (ESS) if competition is reduced, for example by co-evolutionary specializations with plants. The possibility that it has evolved as a mixed ESS (i.e. individuals can both hover and perch depending on the resource level) is discussed. The evolution of optimal foraging strategies is discussed, and it is pointed out that the rate of gain of an animal is independent of the strategy used when all competing foragers use the same strategy, but competitively superior strategies will nevertheless evolve because they are ESSs. Competition between strategies with different energy costs are special, because resource availability determines which strategy is competitively superior. A high-cost strategy can only evolve as a pure ESS at high resource levels, or as a mixed ESS at intermediate levels.     

Evaluation of Rice Cultivars under Greenhouse Conditions for Adult-Plant Resistance to Pyricularia oryzae

Six rice cultivars showing various types of resistance or susceptibility to Pyricularia oryzae in the field were compared under controlled environmental conditions. The resistance of the cultivars with adult-plant resistance was race-specificat early growth stage. On all cultivars tested, blast infection became increasingly reduced on either leaves of adult plants or older leaves, as observed in rice plants of different leaf stages infected with different individual races. Their increase inhost resistance was marked by an apparent transition in infection type and reduced blast severity depending on leaf age and developmental stage of plants. The ranking of disease severity of the adult-plantresistant cultivars to different races was constant during plant development, whereas that of the susceptible cultivars was differential. It is suggested that different degrees of resistance in rice cultivars may exist in seedlings and be consistently maintained during plant development, probably becoming more distinct as rice plants mature.     

南京地区落叶栎林木本植物叶物候研究

叶物候参数长期以来被认为与植物的碳获取的最大化有关,能反映物种的资源利用策略。温带地区因为寒冷冬天的限制,延长叶寿命成为一些物种进行生长发育和繁衍的基础。为探讨叶寿命延长的可能途径(早出叶、晚落叶,或两者兼有),该研究以南京地区两个落叶栎(Quercus spp.)林为研究对象,观测了其中木本植物的出叶物候、落叶物候,并分析了它们与叶寿命之间的关系。结果发现:1)不同物种的出叶开始时间相差较大,出叶早的物种早结束出叶过程;2)不同物种的落叶开始时间相差较大,早开始落叶的物种,落叶持续时间较长,落叶结束时间则相对集中。3)相关分析和回归分析都表明,叶寿命与出叶时间和落叶时间显著关联,但早出叶对叶寿命的延长可能更为重要,因为早出叶相对于晚落叶在物种资源利用上比较具有优势。4)不同物种的出叶时间和落叶时间没有显著相关,可能因为出叶过程和落叶过程是由不同的启动因子引起。这说明延长叶寿命不一定同时通过早出叶和晚落叶来达到。     

Seasonal variation in native hydraulic conductivity between two deciduous oak species

植物抗旱性机制包括耐旱性和避旱性。耐旱性的特征主要是木质部易栓塞性和膨压损失点,而避旱性的特征则是叶片脱落和落叶性。虽然这些机制可以权衡物种连续性,但对于干旱引发的叶片脱落和持续时间的变化如何影响木质部和叶片耐受性之间的关系却知之甚少。在本研究中,我们研究两种具有不同叶片脱落叶行为的橡树(栎属)的耐旱性程度差异。我们预测了Quercus deserticola 在旱季结束时落叶(晚落叶)而面临更大的空化风险,导致其比Q. laeta更耐旱,对水分的利用也更加保守。Q. laeta 只在旱季中期很短的时间内落叶(短落叶)。本研究在墨西哥中部进行,两种橡树各具有单个种群,它们之间相距1.58公里,海拔差为191米。Q. deserticola (晚落叶)多出现在下坡,而Q. laeta (短落叶)多出现在沿坡度海拔较高的地方。我们评估了样地内茎干的水分传导率的季节差异(雨季和旱季),并检测了两个物种间木质部对空化的脆弱性、叶片水分利用和叶片膨松损失点的差异。两种橡树在耐旱性状方面没有显著差异,主要包括木质部易栓塞性、叶片膨松损失点和气孔导度。然而,两种植物在旱季都有不同的表现：短落叶植物对木质部功能的负面影响要比晚落叶植物低。综上所述,两种橡树在植物生理性能方面的季节变化取决于冠层叶面积的减少。     

Rapid accumulation of trihydroxy oxylipins and resistance to the bean rust pathogen Uromyces fabae following wounding in Vicia faba

BACKGROUND AND AIMS: Insect damage to plants leads to wound-activated responses directed to healing of damaged tissues, as well as activation of defences to prevent further insect damage. Negative cross-talk exists between the jasmonic acid-based signalling system that is activated upon insect attack and the salicylic acid-based system frequently activated following pathogen infection. Thus, insect attack may compromise the ability of the plant to defend itself against pathogens and vice versa. However, insect herbivory and mechanical wounding have been shown to reduce fungal infections on some plants, although the underlying mechanisms remain to be defined. This work examines the effects of mechanical wounding on rust infection both locally and systemically in the broad bean, Vicia faba and follows changes in oxylipins in wounded leaves and unwounded leaves on wounded plants. METHODS: The lamina of first leaves was wounded by crushing with forceps, and first and second leaves were then inoculated, separately, with the rust Uromyces fabae at various times over a 24 h period. Wounded first leaves and unwounded second leaves were harvested at intervals over a 24 h period and used for analysis of oxylipin profiles. KEY RESULTS Mechanical wounding of first leaves of broad bean led to significantly reduced rust infection in the wounded first leaf as well as the unwounded second leaf. Increased resistance to infection was induced in plants inoculated with rust just 1 h after wounding and was accompanied by rapid and significant accumulation of jasmonic acid and two trihydroxy oxylipins in both wounded first leaves and unwounded second leaves. The two trihydroxy oxylipins were found to possess antifungal properties, reducing germination of rust spores. CONCLUSIONS: These results demonstrate the rapidity with which resistance to pathogen infection can be induced following wounding and provides a possible mechanism by which pathogen infection might be halted.     

Properties of a mixed ESS candidate in continuous strategy sets

An evolutionarily stable strategy (ESS) is a strategy that if almost all members of the population adopt, then this population cannot be invaded by any mutant strategy. An ESS is not necessarily a possible end point of the evolutionary process. Moreover, there are cases where the population evolves towards a strategy that is not an ESS. This paper studies the properties of a unique mixed ESS candidate in a continuous time animal conflict. A member of a group sized three finds itself at risk and needs the assistance of another group member to be saved. In this conflict, a player's strategy is to choose the probability distribution of the interval between the beginning of the game and the moment it assists the player which is at risk. We first assume that a player is only allowed to choose an exponential distribution, and show that in this case the ESS candidate is an attracting ESS; the population will always evolve towards this strategy, and once it is adopted by most members of the population it cannot be invaded by mutant strategies. Then, we extend the strategy sets and allow a player to choose any continuous distribution. We show that although this ESS candidate may no longer be an ESS, under fairly general conditions the population will tend towards it. This is done by characterizing types of strategies that if established in the population, can be invaded by this ESS candidate, and by presenting possible paths of transition from other types of common strategies to this ESS candidate.     

THE PATHOLOGY OF HYPODERMELLA LARICIS ON LARCH,LARIX OCCIDENTALS

A study of the larch needle disease caused by Hypodermella laricis provided information on the mode of infection, the manner of spread, and the relationship between the parasite and its host, Larix occidentalis. Infection by spores occurs in early spring as soon as the leaves emerge, but there is no evidence that hyphae invade the dwarf shoots. The fungus disrupts the normal abscission mechanism, and the fructifications and spores of H. laricis develop on the leaves which remain attached to the dwarf shoots. The time of precipitation is a decisive factor in the initiation of the disease, i.e., infection occurs only when precipitation coincides with the emergence of young leaves. However, vulnerability to infection diminishes rapidly as the leaves mature. By observing leaf size and noting the time of rainfall, the severity of infection can be accurately predicted. Several adaptive mechanisms related to successful parasitism in H. laricis are discussed.     

Iron Enhances Aluminum-induced Leaf Necrosis and Plant Growth Inhibition in Eucalyptus camaldulensis

The combined effects of excess Fe and Al on Eucalyptus camaldulensis Dehnh. were studied by investigating time course and visible symptoms of leaf necrosis, plant biomass, the status of some antioxidants and pigments and nutrient concentrations. Seedlings were grown hydroponically in nutrient solutions containing 0 or 500 μM AlCl₃, each with a FeSO₄ range of 1, 12 and 120 μM at pH 4.2. Leaf necrosis and plant growth inhibition were induced by Al and enhanced by the increase in Fe concentration. The process from the first appearance of necrotic spots to leaf death (shedding) of a leaf proceeded from a few days to about 20 days after the leaf had fully expanded. Either 120 μM Fe without Al or Al reduced plant growth to a similar extent but 120 μM Fe without Al did not cause leaf necrosis. In leaves, excess Fe (12 and 120 μM) without Al reduced concentration of ASC and GSH, while concentration of Fe, DHA and GSSG and DHA:ASC and GSSG:GSH ratios tended to increase with the increase in Fe concentration in treatment solution with or without Al. At 1 μM Fe, Al increased concentration of DHA and DHA:ASC and GSSG:GSH ratios. Catalase activity in leaves reduced with the increase in leaf Al concentration. At 1 μM Fe, Al greatly reduced concentrations of Fe and chlorophylls in leaves but increase two times Fe concentration in stems. These suggest that the enhancement effects of Fe on Al-induced leaf necrosis and plant growth inhibition can be discussed in context of the excess Fe itself weakens antioxidant capability of ASC–GSH cycle in leaves and greatly reduces plant growth; and the increase in Fe accumulation in stems is involved in Al-induced leaf chlorosis.     

Host availability and the evolution of parasite life-history strategies

Parasites exploit an inherently patchy resource, their hosts, which are discrete entities that may only be available for infection within a relatively short time window. However, there has been little consideration of how heterogeneities in host availability may affect the phenotypic or genotypic composition of parasite populations or how parasites may evolve to cope with them. Here we conduct a selection experiment involving an entomopathogenic nematode (Steinernema feltiae) and show for the first time that the infection rate of a parasite can evolve rapidly to maximize the chances of infecting within an environment characterized by the rate of host availability. Furthermore, we show that the parasite's infection rate trades off with other fitness traits, such as fecundity and survival. Crucially, the outcome of competition between strains with different infection strategies depends on the rate of host availability; frequently available hosts favor "fast" infecting nematodes, whereas infrequently available hosts favor "slow" infecting nematodes. A simple evolutionarily stable strategy (ESS) analysis based on classic epidemiological models fails to capture this behavior, predicting instead that the fastest infecting phenotype should always dominate. However, a novel model incorporating more realistic, discrete bouts of host availability shows that strain coexistence is highly likely. Our results demonstrate that heterogeneities in host availability play a key role in the evolution of parasite life-history traits and in the maintenance of phenotypic variability. Parasite life-history strategies are likely to evolve rapidly in response to changes in host availability induced by disease management programs or by natural dynamics in host abundance. Incorporating parasite evolution in response to host availability would therefore enhance the predictive ability of current epidemiological models of infectious disease.     

Interaction of bean leaf pubescence with rust urediniospore deposition and subsequent infection density

The role of dense straight trichomes on bean leaves associated with race-nonspecific resistance to rust was studied using a settling tower method for uniform inoculation. Abaxial leaf trichomes, which occur only on adult leaves, trapped significant numbers of urediniospores, especially those deposited in clusters, and resulted in reduced effective inoculum reaching the leaf surface. In the field where infection conditions are often less than optimum, leaf pubescence would partially explain the observed resistance. However, the observed increase in total number of spores as well as similar infection densities on fully expanded leaves of glabrous and pubescent genotypes cannot be explained by the trichomes. Other physical or chemical features of the leaf surface such as waxes or cations may play a role. The ecological fit of leaf pubescence is discussed.     

Similarity of epiphyte biomass distribution on Posidonia and artificial seagrass leaves

Epiphyte biomass on Posidonia australis Hook f. and artificial seagrass leaves was analysed in relation to distance from leaf tip at different time periods following the autumnal shedding. The distribution pattern indicated no significant difference in the exponential nature of biomass increase.Similarities between epiphyte biomass distribution on real and artificial leaves is consistent with the hypothesis that biomass distribution on real leaves is not influenced significantly by metabolic interactions between host plant and epiphytes. Although a time component is involved in epiphyte biomass accumulation, the major factor in determining epiphyte biomass distribution is the relative position along the leaf.     

Relative growth rate of leaf biomass and leaf nitrogen content in several mediterranean woody species

In many plant species, herbivory is a major determinant of leaf mortality and it can cause a strong reduction in productive potential. Most predation occurs on young, expanding leaves. Thus, a rapid growth of the leaves can reduce the impact of predation. Furthermore, in cold Mediterranean climates the length of the growing season is constrained to a short period in spring and early summer owing both to low winter temperatures and drought stress in early summer. Therefore, a rapid deployment of leaf area and a high photosynthetic capacity during the spring and early summer might have important positive effects on the final carbon balance of the leaf population. Relative growth rates (RGR) of leaf biomass were measured in 19 woody species typical of Central Western Spain with deciduous and evergreen habits. Highly significant differences were detected in the leaf growth rate of the different species. The differences between species, however, did not correlate either with the mean leaf life-span of each of the species or with other leaf traits such as photosynthetic capacity, specific leaf area or nitrogen content. Leaf growth rate was positively correlated with time elapsed between leaf initiation and fruit maturation, so that species with fruit dispersal in spring and early summer in general had lower leaf growth rates than species with autumn fruit shedding. This relationship shows the effects of the concurrence between vegetative and reproductive organs for nutrients and other resources. Nitrogen concentration in the leaves was very high at the time of bud break, and declined during leaf expansion owing to the dilution associated with the increase in structural components. The rate of nitrogen dilution was, thus, positively related to the leaf growth rate. Relative growth rates calculated for nitrogen mass in leaves were very low compared to the growth in total mass. This suggests that most leaf nitrogen is translocated from the plant stores to the leaf biomass before the start of leaf expansion and that the contribution of root uptake during leaf expansion is comparatively low.     

Studies on the influence of leaf washings on infection by Mycosphaerella ligulicola

Infection of chrysanthemum leaf discs by spores of Mycosphaerella ligulicola was increased in the presence of concentrated leaf washings obtained from five plant species tested. On separation of leaf washings by dialysis the residue stimulated infection and caused aggregation of hyphae on leaf and cellophane surfaces. In the presence of the diffusible fraction an extensive branching network of hyphae developed but there was no increase in infection except in the presence of the chrysanthemum diffusate. Infection of leaf discs of certain plants normally resistant to M. ligulicola occurred in the presence of concentrated leaf washings. Leaf washings from a resistant variety of chrysanthemum had the same effects as those from a susceptible variety. An extract of chrysanthemum leaves was less effective in stimulating infection than was a leaf washing. Defined nutrient media and potato extract stimulated infection but caused the development of different growth habits on leaf surfaces. The active components of the dialysis residue of chrysanthemum leaf washings were heat-stable but apparently volatile. Eluates from strips of certain regions of paper chromatographs of leaf washings and potato extract caused increased infection together with development of characteristic growth habits of the fungus on the leaf.     

Nitrate availability and shoot area development in small willow (Salix viminalis)

Abstract. The effect of nitrate supply upon leaf area development in willow ( Salix viminalis ) was investigated. Rooted cuttings were grown in culture solution at two different rates of exponentially-increasing nitrate supply and with free access to other nutrients. During the lag-phase in attaining stable nutrition, the rate of production of unfolded leaves, in the treatment with poorer N-supply, decreased. After acclimation to the different N-availabilities, the rate of dry matter increase per plant nitrogen was constant and the same in both treatments. The ratio of total leaf area to total amount of nitrogen in the plant was slightly higher in the treatment with poorer nitrogen supply. During the exponential (stable) phases of growth, nearly all the increase in total leaf area was attributable to main-stem leaves; large numbers of leaves on axillary shoots in the treatment with better N-supply contributed little to the total area. During the stable growth phase, the rate of production of unfolded leaves was the same in both treatments. Higher maximum values of relative rate of increase in area of single leaves were found with a better N-supply and at higher leaf positions on the main stem. The duration of leaf expansion was not much affected by N-supply. Final leaf size was greater at higher positions on the main-stem. Within a treatment (investigated for poorer N-supply), differences in final leaf size were proportional to differences in numbers of epidermal cells. Final leaf size was greater, at the same node, in the treatment with better N-supply. Better N-availability had increased the final size but not the number of epidermal cells. It was concluded that the higher exponential rate of total area increase with a better N-supply was largely associated with higher rates of expansion in epidermal cells.     

The Translocation of 14C-Labelled Assimilates in Tomato Plants Infected with Alternaria Solani (Ell. and Mart.) Jones and Grout

A quantitative study was made of the effect of infection bya facultative parasite, Alternaria solani, on the translocationof ¹⁴C-labelled assimilates in the tomato plant. In a plantwith a single diseased leaf, this leaf retained a greater proportionof its assimilated radiocarbon during the early stages of infection.The export of assimilates from a diseased leaf when diseasedevelopment was purposely retarded and was found to be similarto that from a control leaf. At a late stage in the disease,where there was extensive chlorosis and necrosis, export fromdiseased leaves was generally increased and the distributionof the translocated assimilates was altered. Non-infected leaveswere at first unaffected by the disease, but later there wasincreased contribution to the actively growing regions of theplant. In some cases there was a slight increase in total exportfrom healthy leaves but, more generally, a change in the distributionof assimilates occurred. In particular, there was a consistentincrease in activity in the roots. This response to infectionwas not related to the amount of disease present. No evidencewas obtained for an increased import into infected leaves fromnon-infected leaves at any stage of the disease.     

Does leaf shedding increase the whole-plant carbon gain despite some nitrogen being lost with shedding?

When old leaves are shed, part of the nitrogen in the leaf is retranslocated to new leaves. This retranslocation will increase the whole-plant carbon gain when daily C gain : leaf N ratio (daily photosynthetic N-use efficiency, NUE) in the old leaf, expressed as a fraction of NUE in the new leaf, becomes lower than the fraction of leaf N that is resorbed before shedding (R(N)). We examined whether plants shed their leaves to increase the whole-plant C gain in accord with this criterion in a dense stand of an annual herb, Xanthium canadense, grown under high (HN) and low (LN) nitrogen availability. The NUE of a leaf at shedding expressed as a fraction of NUE in a new leaf was nearly equal to the R(N) in the LN stand, but significantly lower than the R(N) in the HN stand. Thus shedding of old leaves occurred as expected in the LN stand, whereas in the HN stand, shedding occurred later than expected. Sensitivity analyses showed that the decline in NUE of a leaf resulted primarily from a reduction in irradiance in the HN stand. On the other hand, it resulted from a reduction in irradiance and also in light-saturated photosynthesis : leaf N content ratio (potential photosynthetic NUE) in the LN stand.     

Whole-tree water use efficiency is decreased by ambient ozone and not affected by O3-induced stomatal sluggishness

Steady-state and dynamic gas exchange responses to ozone visible injury were investigated in an ozone-sensitive poplar clone under field conditions. The results were translated into whole tree water loss and carbon assimilation by comparing trees exposed to ambient ozone and trees treated with the ozone-protectant ethylenediurea (EDU). Steady-state stomatal conductance and photosynthesis linearly decreased with increasing ozone visible injury. Dynamic responses simulated by severing of a leaf revealed that stomatal sluggishness increased until a threshold of 5% injury and was then fairly constant. Sluggishness resulted from longer time to respond to the closing signal and slower rate of closing. Changes in photosynthesis were driven by the dynamics of stomata. Whole-tree carbon assimilation and water loss were lower in trees exposed to ambient O(3) than in trees protected by EDU, both under steady-state and dynamic conditions. Although stomatal sluggishness is expected to increase water loss, lower stomatal conductance and premature leaf shedding of injured leaves aggravated O(3) effects on whole tree carbon gain, while compensating for water loss. On average, WUE of trees exposed to ambient ozone was 2-4% lower than that of EDU-protected control trees in September and 6-8% lower in October.