The life-cycle and host preferences of the bird cherry-oat aphid, Rhopalosiphum padi L., and their bearing on the theories of host alternation in aphids

Emigrants of the host-alternating bird cherry-oat aphid fly from the primary host to various grasses before the end of June, even in the absence of natural enemies on bird cherry. They fly before the leaves of bird cherry mature and at a time when numbers of insect predators are rapidly increasing. In the autumn gynoparae and males return to bird cherry whose senescing leaves again provide the aphids with a rich source of food. The growth efficiency of R. padi on bird cherry and oats indicates that the quality of the available food on the two hosts is markedly different.
Apterous exules prefer oats to young bird cherry leaves and survive best on oats. Gynoparae prefer bird cherry leaves and only successfully produce offspring on the primary host. This, and other work published on R. padi , support the premise that a change in the aphid's preferences causes the change of host.
The dual-discrimination theory of host alternation in aphids is examined in the light of this and other published work.     

Feeding substrates and behaviors of Western cherry fruit fly (Diptera: Tephritidae)

A study was conducted to determine the abundance of potential foods and the feeding substrates and behaviors of the western cherry fruit fly, Rhagoletis indifferens Curran (Diptera: Tephritidae), in 2005, 2006, and 2007 in central Washington state. Aphid colonies with honeydew, a presumed food source for flies, were not seen on randomly selected branches of sweet cherry trees, Prunus avium L., but leaves with cherry juice, fruit that were damaged, and leaves with bird feces were commonly seen, especially later in the season. Grazing, a behavior in which the mouthparts rapidly move up and down and touch plant surfaces without discrete substances visible to the human eye, was seen more frequently in flies on leaves than on fruit. Grazing occurred more frequently than feeding on extrafloral nectaries (EFNs) on leaf petioles, cherry juice on leaves, and bird feces on leaves. The percentages of females and males that grazed on leaves were not different in 2 of 3 yr, but the percentage of females that grazed was higher in a third year. Percentages of female and male flies that fed on EFNs, cherry juice, and bird feces did not differ. More flies grazed the tops than bottoms of leaves. Flies also grazed on leaves of apple, pear, and grape. The results support the hypotheses that R. indifferens feeds mostly on leaves rather than fruit and that leaf surfaces may be the main feeding substrates for R. indifferens throughout the season.     

Chemical aspects of host‐acceptance behaviour in the bird cherry‐oat aphid Rhopalosiphum padi: host‐acceptance signals used by different morphs with the same genotype

Host acceptance by gynoparae and winged virginoparae of the bird cherry‐oat aphid Rhopalosiphum padi (L.) is investigated utilizing leaves and aqueous extracts of the primary and secondary hosts, as well as nonhost plants. Gynoparae are specialized to reproduce on bird cherry Prunus padus L., whereas virginoparae reproduce and feed on various grasses. Host acceptance is assessed using levels of reproduction and survival for adults, as well as survival for nymphs. Little is known of host acceptance by nymphs. The data show that gynoparae and winged virginoparae produce nymphs almost exclusively on their host plants, bird cherry and barley leaves, respectively, over 72 h. When tested with aqueous plant extracts, however, gynoparae produce nymphs almost exclusively on bird cherry extract and progeny numbers are found to be similar to those on plant leaves. Few nymphs are produced on artificial diet. By contrast, winged virginoparae produce nymphs on aqueous extracts of barley, bird cherry and bean, as well as on artificial diet. The numbers of nymphs deposited by gynoparae are similar on aqueous extracts of bird cherry leaves collected at different times during the growing season. When extracts from leaves of various Prunus species are tested, only leaves of P. padus and Prunus virginiana stimulate parturition. Oviparae, the sexual female nymphs of gynoparae, survive well for 96 h on both bird cherry and barley leaves but not on bean seedlings, whereas nymphs of winged virginoparae survive well only on barley leaves. They do not survive for 96 h on any plant‐leaf extracts, although they do survive on artificial diet.     

Probing behavior of bird cherry-oat aphid Rhopalosiphum padi (L.) on native bird cherry Prunus padus L. and alien invasive black cherry Prunus serotina Erhr. in Europe and the role of cyanogenic glycosides

The probing behavior of bird cherry-oat aphid Rhopalosiphum padi was studied on its natural winter host in Europe, the bird cherry Prunus padus, and on the invasive black cherry Prunus serotina, on which spring generations of R. padi do not survive. The EPG-recorded behavior of R. padi on bird cherry and black cherry showed differences in crucial aspects of probing and feeding. The period of the pre-phloem penetration was twice as long and rarely interrupted in aphids on bird cherry as opposed to aphids on black cherry. On black cherry, there was a considerable delay between finding and accepting the phloem. Aphids that had sampled phloem sap either refused to ingest it or the ingestion periods were very short. Amygdalin and prunasin (cyanogenic glycosides present in leaves of Prunus) seriously impeded ingestion activities when applied in pure sucrose diet. The role of amygdalin and prunasin in winter host plant selection and host alternation in R. padi is discussed.     

Induced proteolysis within the bird cherry leaves evoked by Rhopalosiphum padi L. (Hemiptera, Aphidoidea)

The objectives of this study were to elucidate the impact of bird cherry-oat aphid (Rhopalosiphum padi L.) feeding on functioning of the proteolytic machinery in bird cherry leaves. Biochemical analyses proved that R. padi feeding in tissues of primary host stimulated activity of the two major fractions of proteinases (extracted at the optimal pH values: 5.0 and 7.0). Additionally, it has been demonstrated that aphids' feeding on bird cherry led to a decline in levels of albumins and globulins (main protein fractions in P. padus leaves). The opposite tendency, regarding the amounts of these protein fractions was ascertained at the phase of disappearance of R. padi population on tested shoots. Furthermore, it is reported that an increase in activity of the analysed enzymes and a decline in the content of tested protein fractions, were proportional to density of aphid individuals developing on P. padus side shoots. It is hypothesized that long-term R. padi feeding may lead to intensifying the catabolic processing of proteins by the activated proteolytic machinery in bird cherry leaves. The multi-level biological functions of endogenous plant proteinases and their significance in triggering the defense reactions in aphid-infested plant tissues are discussed.     

Reproduction and probing behaviour of the bird cherry‐oat aphid Rhopalosiphum padi on detached leaves and leaves on intact seedlings of the winter host,bird cherry Prunus padus

Video recording and electrical penetration graph techniques are used simultaneously to investigate host acceptance (in terms of reproduction) and probing activities of autumn migrants (gynoparae) of the bird cherry‐oat aphid Rhopalosiphum padi (L.) on detached leaves and on leaves on intact plants of the winter host, bird cherry Prunus padus. There are no significant differences between the times taken to first parturition or the number of nymphs produced over a 6‐ or 18‐h period on intact plants or detached leaves. Stylet probing activities (i.e. total probe duration, xylem ingestion, the time to first phloem contact and phloem salivation) are also similar in the two situations.     

Effects of leaf age on feeding 'preference' and egg laying in the chrysomelid beetle, HaZtica lythri

Final instar larvae and adults of the chrysomelid beetle Haltica lythri Aubé were tested for their feeding preferences on young and senescent leaves of their host plant Epilobium hirsutum L., and found to spend most of the time feeding on young leaves when in a free-choice situation. Females fed senescent leaves laid significantly fewer eggs than females fed young leaves. Feeding on senescent leaves is discussed in an ecological context.     

Responses of holocyclic and anholocyclic Rhopalosiphum padi populations to low‐temperature and short‐photoperiod induction

The different life cycles of aphid species make these organisms good models for studying the short‐term consequences of sex. The bird cherry‐oat aphid Rhopalosiphum padi has a wide geographic distribution and correspondingly different life cycles. In this study, the life cycles of R. padi collected from six different regions in China were characterized experimentally by comparing the responses of holocyclic and anholocyclic populations to low‐temperature and short‐photoperiod induction. Clones collected from Chuzhou, Taian, and Taigu consistently reproduced via obligate parthenogenesis, whereas clones from Hami and Baicheng were holocyclic in their response, and those from Lanzhou were both holocyclic and anholocyclic. Prolonged exposure to low temperature and a short photoperiod (LS) had negative effects on the offspring of anholocyclic aphids with regard to adult lifespan, total longevity, and fecundity compared with aphids maintained at a normal temperature and a long photoperiod (NL). Holocyclic LS R. padi had longer developmental times at all nymph stages, a shorter adult lifespan, shorter total longevity, and a lower fecundity than NL counterparts. The adult prereproduction period of gynoparae was significantly longer than that of virginoparae, and the total longevity of gynoparae was significantly shorter than that of virginoparae. Moreover, the net reproductive and gross reproduction rates, as well as the total fecundity, were roughly fivefold higher in virginoparae than in gynoparae, indicating that there is the short‐term cost of sex. When maintained on their secondary host (Triticum aestivum), gynoparae, males, and oviparae produced by holocyclic populations could survive, and gynoparae produced oviparae. However, under NL conditions, oviparae could not produce overwintering eggs on the secondary host, whereas a few overwintering eggs were generated by oviparae under LS conditions. Taken together, these results illuminate the complexity of insect responses and contribute to a complete understanding of the aphid life cycle and its evolution.     

Reproduction site preference and performance by sexuparae,and mating behavior of their sexual generation on the primary host plant in a heteroecious aphid,Neothoracaphis yanonis (Homoptera: Aphididae)

We studied reproduction site preference and performance by the sexuparae (autumnal migrants) of Neothoracaphis yanonis and mating behavior of their sexual generation on its primary host plant, Distylium racemosum. The sexuparae preferred younger leaves of D. racemosum for settlement and imbibing leaf sap, and they produced more offspring there than on older leaves. Thus, it is suggested that the sexuparae selected more nutritious younger leaves to increase their own fecundity. The offspring consist of yellowish dark-grey and creamy yellow type nymphs, which develop into small males and large oviparae, respectively. Yellowish dark-grey nymphs were deposited gregariously on the basal part of the abaxial surface of a leaf blade, while creamy yellow nymphs were deposited evenly over the abaxial surface. Such a localized distribution pattern probably resulted from sexuparae's strategy to enhance a possibility for their male offspring to mate with oviparae at the base of leaf blade, over which oviparae crawl to twigs. During mating and sometimes afterwards, the ovipara incidentally carried a male on her back and crawled downward on twig to find an axillary bud for oviposition. This behavior may be advantageous to males, who can guard their mate during this period against other conspecific males. Such mate-guarding behavior seems to be related to the development of dual mate-seeking strategies, in which males try to copulate, first at the basal part of midrib, and second on the axillary bud.     

Influence of urban tributaries on freshwater mussel populations in a biologically diverse piedmont (USA) stream

We investigated the distribution patterns of senescent and green leaf litter patches on a streambed to evaluate the hypothesis that the different immersion times of senescent leaves in long-term benthic storage and newly retained green leaves affect streambed distribution patterns in summer (June, July, and August). We counted all the leaf litter patches in the streambed of the study reach, comparing the physical condition of patches and classifying the trapping obstacles associated with each patch type. The distribution patterns of senescent and green leaf litter patches differed. Green leaf litter patches were more numerous at every sampling date, with most trapped by cobbles, whereas senescent leaf litter patches were retained by twig obstacles, backwaters, and cobbles. In June and July, senescent leaf litter patches were located in significantly deeper stream areas than were green leaf litter patches. The distribution of senescent leaves would be primarily determined during spring snowmelt-driven floods. We speculate that senescent leaves were originally located at the edges of pools in the main flow pathway of the channel, which overflowed in the floods. We conducted flume experiments to clarify the transport characteristics of senescent and green painted maple and manchurian alder leaves in the water column. Our hypothesis was that the transport characteristics of each leaf type differ when they first enter the water, because of differing leaf properties. The flume experiments showed that duration of surface flotation differed for senescent and green leaves and for the two tree species. These differences in the duration of leaves on the streambed and in the floating time of green leaves of different trees ensure varied food resources for macroinvertebrates in various physical conditions. Handling editor: B. Oertli     

Green leaves enhance the growth and development of a stream macroinvertebrate shredder when senescent leaves are available

1. Freshly fallen green leaves and flowers of terrestrial plants enter temperate streams in spring and summer, when senescent leaf litter is often scarce. These resources appear to provide good supplementary food for macroinvertebrate shredders, but have some potential shortcomings as food or case material for caddisflies. 2. To compare suitability of green leaves or flowers and senescent leaves for the growth and development of stream shredders, we reared the caddisfly Lepidostoma complicatum in the laboratory with treatments that provided larvae with senescent (oak) and green (oak or maple) leaves separately, and also together, in case the combined use of both types of leaf may benefit the shredder. 3. Larvae supplied with green leaves alone grew at 65% of the rate of those provided with senescent leaves alone, due to their lower consumption rate. No individuals given green leaves alone developed into adults, whereas 70% of the individuals given senescent leaves alone did. Green leaves may inhibit larval consumption due to their high phenol content, or they may be unsuitable for case material because they are less tough than senescent leaves. 4. Larvae supplied with both senescent and green leaves (or flowers) had a higher growth rate and developed faster, than those given senescent leaves alone, whereas the proportions of successfully emerged individuals did not differ. Lepidostoma probably benefits from the higher nitrogen content of the green leaves when used together with senescent leaves. 5. These results suggest that green leaves (or flowers) cannot serve as an alternative food resource to senescent leaves, but that they can enhance the growth and development of a Lepidostoma stream shredder if senescent leaves are also available.     

Development of the aphid parasitoid,Aphidius ervi, in asexual and sexual females of the pea aphid,Acyrthosiphon pisum, and the blackberry-cereal aphid,Sitobion fragariae

The aphid parasitoid,Aphidius ervi Haliday, overwinters in larval diapause. The possibility that the parasitoid might prefer sexual (oviparae) rather than asexual females (virginoparae) as overwintering hosts (oviparae predominate in autumn when host numbers are generally declining) was tested by comparing these aphid morphs as potential hosts. Two host species were also examined, the pea aphid,Acyrthosiphon pisum (Harris), and the blackberry aphid,Sitobion fragariae (Walker). The parasitoids took longer to develop inS. fragariae than inA. pisum but the development of non-diapausingA. ervi was similar in sexual and asexual females. This observation, together with the greater variation in the duration of the different parasitoid stadia inS. fragariae, indicated that the parasitoid is specialized on the pea aphid. In photophases of 12 h and longer, the proportion ofA. ervi entering diapause inA. pisum oviparae was higher than in virginoparae. The critical daylength (where 50% of parasitoids entered diapause) was therefore longer in oviparae (12.6 h) than in virginoparae (11.7 h) with the inference that parasitoids developing in the oviparae would enter diapause earlier in the field. InS. fragariae, critical day-lengths were similar in both aphid morphs. The duration of diapause was unaffected by host morph and emergence in short days (10:14 L:D) occurred over a long period (c. 60 days).     

Morph determination in the bird cherry-oat aphid, Rhopalosiphum padi L

Both crowding and poor nutrition induce the appearance of emigrants in Rhopalosiphum padi L. No emigrants developed when the aphid was reared in isolation for six successive generations on actively growing leaves of bird cherry. However, emigrants developed as soon as the leaves ceased to grow or when the aphids became crowded. Crowding of mothers and postnatal crowding of the nymphs of apterous exules both influenced the induction of alate exules. The highest proportion of alate exules developed when both mother and offspring were crowded. Short day-length induced the appearance of gynoparae and males. For 50% production of gynoparae and males, over the range of temperature 10–18d?C, a 1.75 h reduction in day-length is required for every 4d? increase in temperature. At 18 d?C, even at day-lengths as short as 6 h, exposure for three generations is required before all offspring become gynoparae or males. Low temperature and short day-length was ineffective in inducing the development of gynoparae or males in the first three generations developing from the fundatrix.     

Shredder colonization and decomposition of green and senescent leaves during summer in a headwater stream in northern Japan

We conducted a decomposition experiment using green and senescent maple and alder leaves in a coastal headwater stream in Hokkaido, northern Japan, during June and July 2000. We estimated whether shredder colonization on the leaves and leaf breakdown differed between green and senescent leaves during the experimental period. Late-instar Lepidostoma complicatum (Trichoptera) and Sternomoera rhyaca (Amphipoda) were the predominant shredder taxa among the macroinvertebrates that colonized litterbags. There was no significant difference in shredder colonization between green and senescent leaves although we found a significant difference between maple and alder leaves. The colonization patterns of large individuals of L. complicatum and S. rhyaca differed from those of small individuals. All decomposition coefficients of green and senescent leaves were high. During the experiment, decomposition was significantly faster in maple than in alder leaves, although no significant difference was found between green and senescent leaves. However, the fragmented nitrogen portion was higher in green leaves than in senescent leaves during the experiment. Higher nitrogen release (2–2.5 times more) as particulate organic matter in green than in senescent leaves indicates that green leaves may be a potentially valuable food resource for other macroinvertebrate collector–gatherer species.     

Translocation and Metabolism of Ricinine in the Castor Bean Plant, Ricinus communis L

Ricinine-3,5-¹⁴C (N-methyl-3-cyano-4-methoxy-2-pyridone) administered to senescent leaves of Ricinus communis L. was translocated to all other tissues of the plant. Developing fruit and especially seeds were found to be labeled the most rapidly. Young growing leaves and other developing tissues of the plant imported ricinine from the senescent leaves much more quickly than mature leaves. Relative intensities of the radioactive ricinine imported and deposited in various tissues indicate a possible functional role of ricinine in the castor bean plant. Data on N-demethyl ricinine presented here may stimulate interest in the possible physiological role of the ricinine to N-demethyl ricinine interconversion.     

Photoinhibition and photoprotection in senescent leaves of field-grown wheat plants

Photosynthesis, photosystem II (PSII) photochemistry, photoinhibition and the xanthophyll cycle in the senescent flag leaves of wheat (Triticum aestivum L.) plants grown in the field were investigated. Compared to the non-senescent leaves, photosynthetic capacity was significantly reduced in senescent flag leaves. The light intensity at which photosynthesis was saturated also declined significantly. The light response curves of PSII photochemistry indicate that a down-regulation of PSII photochemistry occurred in senescent leaves in particular at high light. The maximal efficiency of PSII photochemistry in senescent flag leaves decreased slightly when measured at predawn but substantially at midday, suggesting that PSII function was largely maintained and photoinhibition occurred in senescent leaves when exposed to high light. At midday, PSII efficiency, photochemical quenching and the efficiency of excitation capture by open PSII centers decreased considerably, while non-photochemical quenching increased significantly. Moreover, compared with the values at early morning, a greater decrease in CO₂ assimilation rate was observed at midday in senescent leaves than in control leaves. The levels of antheraxanthin and zeaxanthin via the de-epoxidation of violaxanthin increased in senescent flag leaves from predawn to midday. An increase in the xanthophyll cycle pigments relative to chlorophyll was observed in senescent flag leaves. The results suggest that the xanthophyll cycle was activated in senescent leaves due to the decrease in CO₂ assimilation capacity and the light intensity for saturation of photosynthesis and that the enhanced formation of antheraxanthin and zeaxanthin at high light may play an important role in the dissipation of excess light energy and help to protect photosynthetic apparatus from photodamage. Our results suggest that the well-known function of the xanthophyll cycle to safely dissipate excess excitation energy is also important for maintaining photosynthetic function during leaf senescence.     

Plant quality, progeny sequence, and the sex ratio of the sycamore aphid, Drepanosiphum platanoidis

The reproductive sequence of sexuparae of the sycamore aphid, Drepanosiphum platanoidis (Schrank) (Homoptera: Callaphididae), observed in the field and glasshouse was examined to elucidate the relationship between host nutrition and sex ratio. The observations were made in autumn when the transition from parthenogenetic to sexual reproduction occurred. A switch occurred in the reproductive sequence of the sexuparae. The production of apterous sexual females (oviparae) was followed by alate males, after a brief reproductive pause lasting 2.5–4.0 days. The switchover period was relatively constant across treatments and years. On poor hosts, the first appearance of the oviparae was a day later, whereas the first appearance of males was a day earlier than those on good nutrition. The primary sex ratio (i.e., proportion of males) was numerically female biased. Under semi‐natural field conditions it was 0.22, but altered to 0.18 and 0.29, respectively, under poor and good host nutrition in the glasshouse. In both the field and glasshouse, nymphal development time was significantly longer in oviparae (ca. 22 days) than in males (ca. 18 days). The time from birth to adult was significantly shorter on good hosts (oviparae: 20 days; males: 17 days) than on poor hosts (oviparae: 23 days; males: 19 days). The proportion of nymphs surviving to the adult moult was not dependent on sex in both the field and glasshouse, but the chances of survival were slightly enhanced under good (88%) compared with poor (82%) host nutritive status. In conclusion, the order of morph production and differences in maturation times ensured the synchronisation of the sexes so that mating and oviposition opportunities were optimised before the onset of winter. The female‐bias in the sex ratio maximised the fitness of sexuparae, and reduced the chances of local mate competition by economising in members of the competing sex.     

Chlorophyll a fluorescence, enzyme and antioxidant analyses provide evidence for the operation of alternative electron sinks during leaf senescence in a stay-green mutant of Festuca pratensis

Mutation of the sid gene in Festuca pratensis prevents chlorophyll degradation. The senescing leaves retain their chlorophyll complement and stay green. Nevertheless, CO2 assimilation and ribulose-bisphosphate carboxylase/oxygenase content decline in both mutant and wild-type plants. Photosynthesis and chlorophyll a fluorescence measurements were performed in air and at low oxygen to prevent photorespiration. The maximum extractable activity of ribulose 1,5 bisphosphate carboxylase was higher in the senescent mutant leaves than in those of the wild-type control hut Mas much lower than that observed in the mature leaves of either genotype. The activation state of this enzyme was similar in mutant and wild-type lines at equivalent stages of development. Analysis of chlorophyll a fluorescence quenching with varying irradianco showed similar characteristics for mature leaves of the two genotypes. Genotypic variations in photosystem II (I'SII) efficiency were observed only in the senescent leaves. Photochemical quenching and the quantum efficiency of PSII were greater in the senescent mutant leaves than in (he wild type at a given irradiance. The calculated electron flux through PSII was substantially higher in the mutant with a greater proportion of electrons directed to photorespiration. Maximum catalytic activities of ascorbate peroxidase decreased in senescent compared to mature leaves of both genotypes, while glutathione reductase and monodehydroascorbate reductase were unchanged in both cases. Superoxide dismutase activity was approximately doubled and dehydroascorbate reductase activity was three times higher in senescent leaves compared with the mature leaves of both genotypes. In no case was there a difference in enzyme activities between mutant and wild type at equivalent growth stages. The pool of reduced ascorbate was similar in the mature leaves of the two genotypes, whereas it was significantly higher in the senescent leaves of the mutant compared with the wild type. Conversely, the hydrogen peroxide content was significantly higher in the mature leaves of the wild type than in those of the mutant, but in senescent leaves similar values were obtained. In leaves subjected to chilling stress the reduced ascorbate pool was higher in both mature and senescent leaves of the mutant than in their wild-type counterparts. Similarly, the hydrogen peroxide pool was significantly lower in both mature and senescent leaves of the mutant than in the wild type. We conclude that, in spite of deceased CO₂ assimilation, the mutant is capable of high rates of electron Slow. The high ascorbate/hydrogen peroxide ratio observed in the mutant, particularly at low temperatures, suggests that the senescent leaves are not subject to enhanced oxidative stress.     

Cereal aphid responses to sex pheromones and host-plant odours in the laboratory

Abstract. The movement of Sitobion fragariae (Wlk.) (Homoptera: Aphididae) males towards odours from conspecific sexual females (oviparae) and the aphid sex pheromone component (4a S ,7 S ,7a R )-nepetalactone were demonstrated in a linear-track olfactometer. Males of the closely related Sitobion avenae (Fab.) were attracted to odours from conspecific oviparae and also oviparae of S. fragariae. These results were consistent with the recent identification of this nepetalactone isomer as the major component of the sex pheromone in these species. Males of S.fragariae were not significantly attracted to oviparae of S.avenae. indicating some qualitative or quantitative difference in the pheromones of the two species. Males did not respond to the related (1R,4aS,7S,7aR)-nepetalactol or to host-plant odours. Gynoparae of these species moved towards the nepetalactone and also to host-plant odours. A combination of both stimuli was more attractive than plant odour alone. Gynoparae of S.fragariae also responded significantly to the combined treatment when this was tested against the nepetalactone.     

The effect of 6-benzylaminopurine (benzyladenine) on senescence and chocolate spot (Botrytis fabae) of winter beans (Viciafaba)

In laboratory trials on detached bean leaves, benzyladenine delayed both senescence and aggressive attack of chocolate spot disease. In field plots, over 80% of the aggressive spots were on senescent leaves and the distribution of aggressively attacked leaves was predictable from the distribution of senescent leaves. The total frequency of aggressively attacked leaves per shoot was apparently related to the frequency of senescent spotted leaves. Although benzyladenine significantly reduced leaf senescence in field plots sprayed in May, the senescent leaves were more frequently attacked aggressively. Wider plant spacing (14 in instead of 8 in) similarly increased the ratio of aggressive attack to senescence, and the individual fields retained until June the differences in infection earlier established by frost damage.