Competition Alters Temporal Dynamics of Sporulation in the Wheat Stem Rust Fungus

To investigate the effects of competition on the timing of pathogen reproduction, urediniospores of two strains of Puccinia graminis f.sp. tritici (SR22 and SR41) were inoculated onto leaves of wheat seedlings singly and in 1 : 1 mixture at three inoculum densities. On randomly sampled leaves, uredinia were counted 9 days after inoculation and urediniospores were collected and quantified every other day from the seventh to the 29th day after inoculation. Increases in inoculum density resulted in progressively smaller increases in uredinial numbers. However, total urediniospore production per leaf was not significantly affected by inoculum, and hence uredinial, density over a range of approximately 10-300 uredinia on the leaf. Total urediniospore production per uredinium generally decreased with increasing inoculum or uredinial density. At high densities, sporulation per uredinium peaked earlier in the sporulation period, had a less distinct peak, and dropped off earlier than for the lower densities. Logistic model fits to cumulative sporulation curves over time revealed that strain SR41 had a greater epidemic rate parameter (r) than SR22 at low and intermediate inoculum or uredinial densities, while SR22 had a higher r-value than SR41 at high density. Both strains also exhibited greater r-values in the presence of the other strain than when alone. Results suggest that strains may have different ecological strategies in their timing of reproduction, and that both intra- and interstrain competition can have complex effects on the temporal dynamics of sporulation in pathogen strains.     

Relative importance of components affecting the leaf rust progress curve in wheat

Summary The relative importance of various components (latent period, uredium area, infectious period, infectibility, sporulation and weather) affecting the progress of leaf rust (Puccinia recondita) in wheat, in the field, was determined by calculating the effect of equivalent changes in the individual components. The effect was calculated in terms of the period of delay in the epidemic reaching 100 per cent severity of leaf rust. Only four components viz., latent period, infectibility, sporulation and weather were found to be important. Further, it was found that these four components were equally important and that they collectively determined the rate of progress of leaf rust in an additive manner.     

Interaction between barley yellow dwarf virus and rust in wheat, barley and oats, and the effects on grain yield and quality

In three separate experiments, the upper leaf surface of the fifth formed leaf of wheat cv. Highbury, the fourth and fifth leaves of barley cv. Julia and the third and fourth leaves of oat cv. Mostyn were inoculated in a spore settling tower with wheat brown rust (Puccinia recondita f. sp. tritici), barley brown rust (P. hordei) or oat crown rust (P. coronata f. sp. avenae), respectively. Fewer pustules developed on distal portions of leaves of plants infected with barley yellow dwarf virus (BYDV) than on similar portions of leaves from virus-free plants. There were no significant differences in the number of pustules on proximal leaf portions. In barley and oats, the number of pustules on distal leaf portions was negatively correlated with the amount of yellowing of the leaf areas scored. In wheat, symptoms of BYDV were mild and leaves were little affected by yellowing. The latent period of rust on wheat and oats was not affected by BYDV. In barley, BYDV reduced the latent period of rust on leaf 5, but not on leaf 4, and reduced it on proximal, but not distal, leaf portions. In other experiments, BYDV reduced the yield of wheat and oats by 44% and 66%, respectively, while BYDV-infected barley was almost sterile. The appropriate rust reduced the yield of wheat, barley and oats by 33%, 13% and 86%, respectively. When infected with both BYDV and rust, yield of wheat and oats was reduced by 63% and 91%, respectively. Neither BYDV nor rust affected the percentage crude protein content of wheat grain, nor did rust affect that of barley. In oats, BYDV and rust each significantly increased crude protein of grain, but rust infection of BYDV-infected plants tended to reduce it.     

Effect of Host Plant Resistance on Haustorium Formation in Cereal Rust Fungi

Haustoria of Puccinia triticina (wheat leaf rust fungus) and P. hordei (barley leaf rust fungus) were isolated from susceptible and partially resistant wheat lines, and susceptible, hypersensitive and partially resistant barley lines. Haustoria were counted and measured. The size of haustoria was similar in the partially resistant and susceptible genotypes but haustoria were smaller in the hypersensitive barley line L94+Pa7. The number of haustoria was reduced in both partially and hypersensitive lines when compared with susceptible ones. Therefore it seems that the reduction in the number of haustoria is a consequence of the resistance that can be attributable either to early abortion of infection units or reduced colony growth. The reduction of the number of haustoria was more pronounced in the adult plant stage.     

Influence of inoculum density of the antagonistic bacteria Pseudomonas fluorescens and Pseudomonas corrugata on sugar beet seedling colonisation and suppression of Pythium damping off

The effect of initial inoculum density of the antagonistic bacterial strains Pseudomonas fluorescens B5 and Pseudomonas corrugata 2140 (10³ to 10⁸ CFU per seed pellet) on sugar beet seedling colonisation, in situ bioluminescence and antagonistic activity towards Pythium ultimum was investigated. Populations of the bacteria colonising sugar beet root systems approached an apparent carrying capacity of 10⁵ to 10⁶ CFU per plant after 12 d growth, irrespective of inoculum density. This meant an up to 320-fold population increase at low inoculum densities and a decrease at high densities. Population densities of both bacteria and their corresponding in situ bioluminescence (resulting from luciferase enzyme expression from the inserted luxAB genes) reached highest levels in the hypocotyl region and in the upper root region 0–20 mm below seed level (10⁴–10⁶ CFU/cm section, 10¹–10³ RLU/cm section) and decreased with root depth. In situ bioluminescence, which indicates physiological activity, was measurable at lowest antagonist initial inoculum density (10³ CFU per seed pellet) and did not increase significantly with increasing inoculum density. Bioluminescence was also significantly correlated with population density. For Pseudomonas fluorescens B5, the total population size per plant and downward colonisation of the root (below 40 mm depth) increased significantly with antagonist inoculum density applied to the seeds. For Pseudomonas corrugata 2140, no significant influence of initial inoculum density on root colonisation was observable. Survival and dry weight of sugar beet seedlings in Pythium infested soil increased significantly with increasing inoculum density of Pseudomonas fluorescens B5, whereas for Pseudomonas corrugata 2140, initial densities of 10⁴ to 10⁶ CFU per seed resulted in maximal survival of plants.     

Heterologous expression of genes mediating enhanced fungal resistance in transgenic wheat.

Three cDNAs encoding the antifungal protein Ag-AFP from the fungus Aspergillus giganteus, a barley class II chitinase and a barley type I RIP, all regulated by the constitutive Ubiquitin1 promoter from maize, were expressed in transgenic wheat. In 17 wheat lines, stable integration and inheritance of one of the three transgenes has been demonstrated over four generations. The formation of powdery mildew (Erysiphe graminis f. sp. tritici) or leaf rust (Puccinia recondita f. sp. tritici) colonies was significantly reduced on leaves from afp or chitinase II- but not from rip I-expressing wheat lines compared with non-transgenic controls. The increased resistance of afp and chitinase II lines was dependent on the dose of fungal spores used for inoculation. Heterologous expression of the fungal afp gene and the barley chitinase II gene in wheat demonstrated that colony formation and, thereby, spreading of two important biotrophic fungal diseases is inhibited approximately 40 to 50% at an inoculum density of 80 to 100 spores per cm2.     

Effect of leaf age and inoculum concentration on the symptoms produced by Gymnosporangiumjuniperi-virginianae on apple

Gymnosporangium juniperi-virginianae, the incitant of cedar apple rust, induced the greatest number of lesions (flecks, pycnial lesions, aecial lesions) on 6- and 8-day-old leaves of single-shoot potted apple plants inoculated and incubated under controlled conditions. The largest pycnial lesions developed on 2- and 4-day-old leaves and the largest aecial lesions on 6-day-old leaves. The number of lesions was positively correlated with the concentration of basidiospores in the inoculum, but lesion size was constant at different inoculum levels. Elevated inoculum concentrations failed to induce pycnia on cv. McIntosh which normally bears only flecks; very low concentrations induced the development of pycnia but not aecia on cv. Rome Beauty, which normally bears aecia. An ‘infection rating’, log₁₀ (10 nd²), where n is the highest number of pycnial lesions per leaf and d is the largest mean diameter of pycnial lesions on any leaf, calculated for four cultivars in greenhouse tests showed a positive correlation with a ‘susceptibility rating’ developed from observations of natural infection of these cultivars in the field.     

Some effects of insecticide deposit patterns on the parasitism of Trialeurodes vaporariorum by Encarsia formosa

The functional response of Encarsia formosa parasitising Trialeurodes vaporariorum on Phaseolus vulgaris was a typical type II functional response (Holling, 1959) where, with increasing host density, the numbers of hosts parasitised by a single parasitoid show a negatively accelerating rise to an upper plateau. A mean maximum parasitism of 6.6 per female per 24 h reached at host densities near 80 larvae per leaf was observed. When presented with insecticide (‘Pynosect 30 ULV’) deposits, the parasite's search pattern changed, conforming to a typical Sigmoid functional response curve. With a uniform cover spray deposit of 27.6 × 10^-7 ml/cm² searching began only at host densities of c. 80 larvae per leaf and the mean maximum parasitism was only 1.5 per female per 24 h. When the insecticide deposit of 27.6 × 10^-7 ml/cm² was in the form of discrete groups of 100 μm droplets, searching began at host densities of c. 40 larvae per leaf but the mean maximum parasitism was low at 2.2. However by using the same discrete deposit pattern but reducing the deposit level to 2.65 × 10^-7 ml/cm² it was possible to obtain the same high mean maximum parasitism as with the untreated control.     

Damage potential of grasshoppers (Orthoptera: Acrididae) on early growth stages of small-grains and canola under subarctic conditions

We characterized the type and extent of grasshopper injury to above- and below-ground plant parts for four crops [barley (Hordeum vulgare L.), oats (Avena sativa L.), wheat (Triticum aestivum L.), and canola (Brassica campestris L.)] commonly grown, or with potential to grow, in central Alaska. Cages were placed on 48 pots containing plants in second to third leaf stages and stocked with 0, 2, 4, and 6 first-instar Melanoplus sanguinipes F. pot(-1). Plants were harvested 22 d after planting. Stem growth of barley and oats was not affected except at the highest grasshopper treatment. In canola, stem biomass was reduced at the medium and high grasshopper treatments, when most of the leaves had been consumed. The highest grasshopper treatment reduced leaf area in barley and oats by approximately 55%, and caused a significant reduction in dry weight of leaves, stems, and roots (41-72%). Wheat and canola plants were smaller than barley and oats across all treatments and, at the highest grasshopper density, above-ground portions of wheat and canola were completely destroyed. Length and surface area of roots of barley and oats were reduced by 20-28% again at the highest grasshopper density, whereas the reduction for wheat and canola ranged from 50 to 90%. There was little or no difference among all grasshopper densities for C-N ratio in leaf and stem tissues of all crops. The results suggest that wheat and canola are more susceptible than barley and oats and that densities > or = 2 pot(-1) (approximately > or = 50 m(-2)) of even very small grasshoppers could cause significant damage in small-grain and oilseed crop production.     

Habitat structure and fish predation: effects on invertebrate colonisation and breakdown of stream leaf packs

1. We investigated the effects of two features of leaf‐pack habitat structure (i.e. mass of a leaf pack and surface area of leaves comprising a leaf pack) and fish predation on colonisation of shredders and leaf breakdown rates in a coldwater stream. Packs were constructed of red maple (Acer rubrum) leaves. 2. A 2 × 3 × 3 factorial experiment was used to manipulate fish predation (exclusion and control cage), leaf‐pack mass (1, 3 and 5 g dry mass) and leaf surface area (small: approx. 17.9 cm², medium: approx. 34.6 cm², large: approx. 65.6 cm²). Exclusion cages had mesh on all sides, whereas control cages lacked mesh on two sides to provide access to fish. 3. Common shredders were Gammarus pseudolimnaeus, Pycnopsyche and Lepidostoma. Shredder biomass per leaf pack increased with the mass of a leaf pack (P < 0.001), but biomass per unit mass of leaf pack did not differ with leaf‐pack mass (P = 0.506). Shredder densities did not respond to the exclusion of fish (P > 0.7) or leaf surface area (P > 0.7), and interactions among treatment factors were not significant (P > 0.2). 4. Breakdown rates were lower for leaf packs comprised of small leaves (P < 0.001) and leaf packs with high mass (P = 0.001). Excluding fish did not significantly affect leaf breakdown rates (P = 0.293), and interactions among treatment factors were not significant (P > 0.3). Breakdown rates were highest when packs consisted of few leaves (i.e. leaf packs with large leaves and low mass) and were colonised by many shredders. 5. Fish predation was not an important factor controlling shredder densities in leaf packs over the spatiotemporal scale of our experiment. Nevertheless, we found shredder colonisation was proportional to leaf‐pack mass and breakdown rates were affected by leaf‐pack size (i.e. number of leaves in a pack). We suspect that fragmentation is the primary mechanism causing the breakdown rates to be dependent on leaf‐pack size.     

Effects of shading and plant density on leaf growth of Viciafaba

Faba beans (Viciafaba cv. Maris Bead) were grown in the field at densities of 20 and 60 plants m^-2 and under zero, 34 and 56% reduction in incoming solar radiation. Measurements of the area of individual leaves were made every two days. Rates of leaf appearance and the duration of expansion of a leaf were virtually unaffected by the experimental treatments. Leaves required approximately 200 ^oC days (above 1 ^oC) from appearance to full expansion. Shading consistently increased specific leaf area but decreased leaf weight only at the greater shading level. At low density leaves were largest under 34% shade and smallest under 56% shade, but at high density there were no significant differences in leaf size.     

Change of virulence of Septoria nodorurn during passage through barley and wheat

The barley biotype of Septoria nodorum was recovered from isolates of the wheat biotype after 2–6 passages through barley. Shifts in the population of the two biotypes during host passage were determined by isolating single spores after passage and observing changes in symptom expression on both hosts. The number of passages required before the barley biotype was recovered varied with the fungus isolate and barley cultivar. Attempts to recover the wheat biotype from isolates of the barley biotype during several passages through wheat were unsuccessful when 10⁵ to 10⁶ conidia m1^-1 were used as inoculum. However, the wheat biotype was recovered from two isolates of the barley biotype during a single passage when a very high concentration of conidia (>10¹² ml^-1) were used as inoculum. From the results of this study and a previous report by the author, the biotype of S. nodorum on barley which occurs in the southeastern United States appears to be largely restricted to barley.     

Effect of density on spatial distribution,pit formation and pit diameter of Myrmeleon acer Walker, (Neuroptera: Myrmeleontidae): patterns and processes

The effect of increasing population density on the formation of pits, their size and spatial distribution, and on levels of mortality was examined in the antlion Myrmeleon acer Walker. Antlions were kept at densities ranging from 0.4 to 12.8 individuals per 100 cm². The distribution of pits was regular or uniform across all densities, but antlions constructed proportionally fewer and smaller pits as density increased. Mortality through cannibalism was very low and only occurred at densities greater than five individuals per 100 cm². Antlions in artificially crowded situations frequently relocated their pits and when more space became available, individuals became more dispersed with time. Redistribution of this species results from active avoidance of other antlions and sand throwing associated with pit construction and maintenance, rather than any attempt to optimise prey capture per se.     

Proteins in intercellular washing fluid from noninoculated and rust-affected leaves of wheat and barley

Proteins in intercellular washing fluid (IWF) from wheat (Triticum aestivum) and barley (Hordeum vulgare) leaves were separated by two-dimensional isoelectric focusing-polyacrylamide gel electrophoresis and stained with Coomassie brilliant blue (CBB) or silver. Intracellular protein from the cut ends of leaves accounted for only a small proportion of total protein in IWF from wheat leaves. When these were heavily infected with the stem rust fungus (Puccinia graminis f. sp. tritici) and grown at 19°C, four infection-related CBB-stainable proteins were detected in IWF.

To compare IWF proteins from wheat and barley leaves infected with the same pathogen, conditions were established that permitted luxuriant growth of stem rust of wheat in barley (exposure to chloroform before inoculation and maintenance at 25°C thereafter). Under these conditions, at least 10 infection-related silver-stainable proteins were detected in IWF from infected wheat in addition to the more than 50 that were of host origin. The electrophoretic properties of 8 of the infection-related proteins were the same as those of 8 infection-related proteins in IWF from barley.

IWF from wheat and barley grown under these conditions was analyzed for Concanavalin A-binding glycoproteins immobilized on nitrocellulose membrane replicas made from gels. Of the many infection-related glycoproteins that were detected in IWF from stem rust-affected wheat, approximately 20 occupied the same positions as those from stem rust-affected barley. The glycoprotein pattern of IWF prepared from wheat leaves grown at 19°C and infected with the leaf rust fungus (P. recondita f. sp. tritici) was markedly different to that of IWF from the same host infected with the stem rust fungus. We conclude that IWF from rust-affected cereal leaves may be a useful source of surface or extracellular proteins from the parasitic mycelium.

     

Seasonal Variation in Pathogenicity of Phytophthora infestans

During a four year study it was shown that the mean lesion area (M.L.A.) developed on leaflets of different potato cultivars by isolates of P. infestans after artificial inoculation varied periodically during the year. Disease symptoms observed changed from pinpoint necroses to regular sporulating lesions depending on the season and the cultivar. Analysis of variance revealed that differences in lesion size between experiments performed in different seasons were significant. The largest M.L.A. was found in late spring or in autumn but autumnal increase of M.L.A. was usually significantly lower than that in spring. Periodicity of M.L.A. was accompanied by a similar phenomenon in sporulation density of P. infestans isolates cultured in vitro. However, the peaks of sporulation, in comparison with peaks of M.L.A. appeared in different seasons. The highest disease level coincided with a decline in sporulation density of P. infestans cultured in vitro and the decrease of M.L.A. corresponded with an increase in sporulation density per cm² of medium. Variation in M.L.A. found in this study seemed not to be related to variability in plant reaction. In this connection a hypothesis has been proposed that the changes of M.L.A. were due to variation in pathogenicity of the fungus.     

Genome-Dependent Factors in the Development of Leaf Phototrophic Tissues in Diploid and Alloploid Wheat Species

Growth and mesostructure of the photosynthetic apparatus were studied in leaves of ten Triticum L. species. Plants with the A^u genome were shown to develop larger leaf assimilation areas due to expanding areas of individual leaves and an increase in the absolute growth rate. Leaf and mesophyll thickness and mesophyll cell size decreased in the G-genome species. Leaf compactness, which depended on cell size and number per unit leaf area and leaf folding, determined the specific patterns of internal leaf organization in wheat species with diverse genotypes. These patterns did not affect cell plastid-to-cytoplasm ratio as shown by the stable indices of cell surface area/cell volume, cell surface area per chloroplast, and cell volume per chloroplast. The structural indices of leaf phototrophic tissues, mesophyll density, and mesophyll CO₂ conductance in alloploids, as compared to diploid species, depended on both ploidy and genome constitution.     

A major QTL for durable leaf rust resistance widely exploited in durum wheat breeding programs maps on the distal region of chromosome arm 7BL

A recombinant inbred line (RIL) population and a set of advanced lines from multiple crosses were used to investigate the leaf rust (Puccinia triticina Eriks.) resistance carried by the durum wheat cultivar Creso and its derivatives (Colosseo and Plinio). One hundred seventy-six RILs from the cross Colosseo × Lloyd were tested under artificial rust inoculation in the field. The response at the seedling stage was also investigated. A major QTL (QLr.ubo-7B.2) for leaf rust resistance controlling both the seedling and the adult open field based-response was mapped on 7BL, with the favourable allele inherited from Colosseo. QLr.ubo-7B.2 showed R ² and LOD peak values for the area under disease progress curve (AUDPC) equal to 72.9% and 44.5, respectively. The presence and location of QLr.ubo-7B.2 was validated by a linkage disequilibrium-based test using two-year field data of 62 advanced lines from 21 crosses with Creso, Colosseo or Plinio as resistance donors. QLr.ubo-7B.2 maps in a gene-dense region (7BL10-0.78-1.00) carrying several genes/QTLs in wheat and barley for resistance to rusts and other fungal diseases. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Morphological plasticity of wheat and barley roots in response to spatial variation in soil strength

Root systems of individual crop plants may encounter large variations in mechanical impedance to root penetration. Split-root experiments were conducted to compare the effects of spatial variation in soil strength on the morphological plasticity of wheat and barley roots, and its relationship to shoot growth. Plants of spring barley (Hordeum vulgare cv Prisma) and spring wheat (Triticum aestivum cv Alexandria) were grown for 12 days with their seminal roots divided between two halves of a cylinder packed with sandy loam soil. Three treatment combinations were imposed: loose soil where both halves of the cylinder were packed to 1.1 g cm^–3 (penetrometer resistance 0.3 MPa), dense soil where both halves were packed to 1.4 g cm^–3 (penetrometer resistance 1 MPa), and a split-root treatment where one half was packed to 1.1 and the other to 1.4 g cm^–3. In barley, uniform high soil strength restricted the extension of main seminal root axes more than laterals. In the split-root treatment, the length of laterals and the dry weight of main axes and laterals were increased in the loose soil half and reduced in the dense soil half compared with their respective loose and dense-soil controls. No such compensatory adjustments between main axis and laterals and between individual seminal roots were found in wheat. Variation in soil strength had no effect on the density of lateral roots (number per unit main axis length) in either barley or wheat. The nature and extent of wheat root plasticity in response to variation in soil strength was very different from that in response to changes in N-supply in previous experiments. In spite of the compensatory adjustments in growth between individual seminal roots of barley, the growth of barley shoots, as in wheat, was reduced when part of the root system was in compacted soil.     

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.