The performance of Brevicoryne brassicae and Myzus persicae in relation to plant age and leaf amino acids

With increasing plant age, total soluble nitrogen of Brussels sprout plants decreased after a peak at 6–9 weeks. The plants then became less suitable for Brevicoryne brassicae and Myzus persicae (particularly the latter) as shown by estimates of the mean relative growth rate (RGR) of the aphids in two experiments. This growth statistic was, however, only poorly correlated with total soluble N and this lack of correlation was particularly marked on plants less than 9 weeks old. Multiple regression on the leaf amino acid spectrum indicated that much of the variability in aphid RGR could be assigned to relatively few amino acids. The complex of asparagine and glutamine gave a positive coefficient for both aphids, but otherwise different amino acids were involved in the regression for each aphid. Attention is drawn to the presence in each regression of an amino acid with a negative coefficient; the inverse correlation of M. persicae RGR with amino-butyric acid may be particularly important in the aphid/hostplant relationship.

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Zusammenfassung In einem ersten Versuch zeigte sich, daß der Gehalt an löslichem Stickstoff in den Blättern mit dem Alter der Pflanzen zwischen 8 und 20 Wochen ständig abnahm. Die relative Wachstumsrate (RGR) von Brevicoryne brassicae veränderte sich dabei kaum, jedoch waren die einzelnen Werte der RGR und des löslichen Stickstoffgehalts der entsprechenden Pflanzen gut korreliert. Dagegen reagierte Myzus persicae unmittelbarer auf das Pflanzenalter als auf den Stickstoffgehalt und ihre RGR nahm ab, je älter die Pflanzen wurden.Im zweiten Versuch wurden jüngere Pflanzen benützt. Hier stieg der Gehalt an löslichem Stickstoff bis zu einem Höhepunkt in der neunten Woche an. Unabhängig vom Stickstoffgehalt nahm die RGR bei M. persicae wieder ab, je älter die Pflanzen wurden. In diesem Versuch wurde auch der, Aminosäuregehalt der Blätter analysiert. Die Variabilität der RGR beider Blattlausarten war mit 4 Aminosäuren korreliert. Sowohl Asparagin wie Glutamin zusammen waren dabei für beide Blattlausarten wichtig; daneben erschienen Threonin und Phenylalanin in der Korrelationsgleichung für B. brassicae, Methionin und Aminobuttersäure in der für M. persicae. Für M. persicae konnten so 42% der gesamten Variabilität der RGR durch die Korrelationsgleichung erklärt werden; für B. brassicae nur 21%.Von besonderem Interesse ist 1) daß sich 42% der M. persicae-Variabilität mit der Verfügbarkeit von nur 4 Aminosäuren erklären lassen, 2) daß die Korrelation zwischen Blattlaus-Wachstumsrate und Asparagin + Glutamin vielleicht dem oft nachgewiesenen Zusammenhang zwischen Pflanzenempfindlichkeit und löslichem Stickstoff entspricht, 3) daß die Wachstumsraten der beiden Blattlausarten negativ mit je einer Aminosäure korreliert sind: mit Phenylalanin bei B. brassicae und, sehr wichtig, mit Aminobuttersäure bei M. persicae und 4) daß die Resultate bezüglich M. persicae ziemlich gut mit Resultaten bei küstlichen Ernährungsversuchen anderer Autoren übereinstimmen.

     

Resistance to attack by Brevicoryne brassicae among plants of Brussels sprouts

Because they remained almost uncolonized by the cabbage aphid (Brevicoryne brassicae (L.)) throughout the growing season, plants of Brussels sprouts were singled out in each of 4 years, from plots heavily infested with the aphid, as possibly being resistant to attack. Clones of these plants were established from cuttings and tested in a controlled environment by inoculation with B. brassicae and later, in the field, by natural infestation. The tests confirmed that some of the plants were resistant to the aphid, and the most resistant of those from the first year of the work proved at least as resistant as any subsequently found. The resistance was expressed as antibiosis, but in the field host non-preference was also shown by incoming winged aphids. The possibility that biotypes of B. brassicae might exist, to which the resistant sprout clones were not necessarily resistant, was investigated using B. brassicae collected from sprouts from each of several areas in England. Eight sprout clones, seven of which were known to be resistant, and the other susceptible, to B. brassicae from Wellesbourne were tested with these other B. brassicae. The results showed that biotypes of the aphid, with differing abilities to colonize respective sprout clones, existed in each area, and of the seven sprout clones resistant to the Wellesbourne aphid, only one appeared never to be fully susceptible to one or more of the other biotypes of B. brassicae.     

Relative feeding rates of Brevicoryne brassicae and Myzus persicae on Brussels sprout plants in relation to their susceptibility to systemic insecticides

Brevicoryne brassicae and Myzus persicae removed similar quantities of ³²P-labelled material from Brussels sprout leaves whether they fed for 24 or 48 h periods. They also removed similar quantities from untreated leaf disks as from leaf disks treated with a sub-lethal dose of menazon. When a lethal dose was used, the uptake of ³²P by B. brassicae was significantly less than by M. persicae. M. persicae excreted a greater proportion of ³²P label in the honeydew than B. brassicae and a greater proportion of the amount absorbed was lost in the progeny of this aphid than in B. brassicae.
B. brassicae was 6.2 times more susceptible than M. persicae to dimethoate acting systemically. When it was applied topically the aphids were equally susceptible.
Considerable variation in uptake of ³²P occurred between replicates and the factors that could influence this are discussed.     

The effect of leaf excision on the performance of Myzus persicae and Brevicoryne brassicae in relation to the nutrient treatment of the plants

Two groups of brussels sprout plants of one variety given different fertilizer treatments proved ‘resistant’ and ‘susceptible’ to both Myzus persicae and Breicoryrie brassicae, as measured by aphid mean relative growth rate. The hypothesis that leaf disc would fail to reflect differences in ‘nutritionallybased’ plant resistance was confirmed with B.brussicue but not M.persicae. Both aphids grew more slowly on discs than leaves of ‘susceptible’ plants; the reverse tendency showed on ‘resistant’ material. Total:(, amino nitrogen proved poorly correlated with aphid performance. Previously published (van Emden & Bashford, 1971) equations relating individual amino acids to aphid performance gave a better fit to the results, but still failed to account for the reduced performance of the aphids on ‘susceptible’ discs.     

Influence of water stress on the glucosinolate profile of Brassica oleracea var. italica and the performance of Brevicoryne brassicae and Myzus persicae

Drought stress alters the chemical composition of plants, which can influence their tolerance to insect herbivory. To evaluate plant chemical responses to drought stress, broccoli, Brassica oleracea L. var. italica Plenck (Brassicaceae), was grown under well‐watered, drought, and water‐logged conditions. The glucosinolate (GS) levels and the performance of two aphid species, the specialist Brevicoryne brassicae (L.) and the generalist Myzus persicae (Sulzer) (both Hemiptera: Aphididae), in relation to water stress conditions were studied. High Performance Liquid Chromatography analysis showed that water stress changed the levels of GS in broccoli plants. Plants grown for 2 weeks under drought stress were significantly smaller and showed decreased levels of total GS when compared with GS contents of well‐watered plants, whereas water‐logged conditions led to a slight increase in the GS contents. A substantial decrease in indolyl GS was detected in water‐deficient plants, whereas aliphatic GS decreased slightly. Analysis of sugar levels in phloem sap of broccoli plants revealed that plants under water‐logged conditions contained the highest amounts of sugars followed by drought‐stressed and well‐watered plants. The two aphid species responded differently to water stress‐induced changes in their host plants. Significantly larger populations of M. persicae were recorded on plants with a limited water supply than on plants grown under well‐watered or water‐logged conditions. Brevicoryne brassicae was less affected by water stress, and similar population sizes were found on plants that were subject to different treatments. Analysis of covariance showed a significant effect of the plants’ water condition but no significant effect of GS content on the performance of M. persicae. However, the specialist B. brassicae remained unaffected by changes induced under water stress conditions.     

The susceptibility of Myzus persicae and Brevicoryne brassicae to 14C-phorate in artificial diets

Colonies of Myzus persicae and Brevicoryne brassicae were established on artificial diets and the honeydew produced by them collected on aluminium foils. The diets were later exchanged for others containing a range of ¹⁴C-phorate concentrations. The honey-dew produced while feeding on the treated diets was collected separately and mortalities were recorded. The diets, aphids and their honeydew were subsequently analysed. About 1.6% of the doses applied initially were found in the diets at the end of the bio-assay period. Thin-layer chromatography of diets containing LC₅₀ concentrations of insecticides showed that phorate and its metabolites were present in similar proportions. The LC₅₀ values were 15.5 nM and 277 nM phorate equivalents for B. brassicae and M. persicae respectively.     

Comparison of feeding behaviour of two Brassica pests Brevicoryne brassicae and Myzus persicae on wild and cultivated brassica species

Feeding behaviour of the specialist Brassicae aphid, Brevicoryne brassicae (L) (cabbage aphid) and the generalist, Myzus persicae, (Sulzer) (peach potato aphid) was monitored electronically on the susceptible cauliflower, Brassica oleracea var. botrytis cv Newton Seale, and a range of 17 Brassica species, B. carinata, B. juncea, B. nigra, B. macrocarpa, and B. villosa var. drepanensis and cultivated brassica varieties, B. oleracea, B. campestris and B. napus. Aphids, monitored for 10 h on the underside of leaves, performed recognisable feeding behaviour on all brassica species. The main differences in feeding behaviour, between M. persicae and B. brassicae, on the susceptible cauliflower Newton Seale, were fewer probes, shorter times to initially reach the phloem but longer times to establish sustained phloem ingestion and the longer times spent, by M. persicae, in xylem ingestion.Feeding behaviour on the range of brassica species tested indicated that generalist and specialist aphids are influenced differently by the host plant. A longer time spent in xylem ingestion was again the major difference in the feeding behaviour of the two aphids. In addition, rejection of passive phloem ingestion, by M. persicae, was not related so closely to increased time spent in non probing activities, as for B. brassicae. This observation indicates that M. persicae does not generally accept or reject brassica species due to the presence of phagostimulants, such as glucosinolates at the leaf surface or along the stylet pathway, unless the concentration is very high. Differences in feeding strategies employed by generalist and specialist aphids on the same plants are discussed.     

Factors affecting the activity of alatae of the aphids Myzus persicae (Sulzer) and Brevicoryne brassicae (L.)

Apparatus was designed for testing the frequency of flights of aphids under different conditions of temperature, relative humidity, light and pressure. Young aphids (1–4 days after metamorphosis) flew more often and showed less individual variability than older ones; with all ages activity increased for the first few hours under experimental conditions. Starving increased activity for the first 1–2 hr. Aphids used in experiments on a second day flew less frequently than controls which had remained on the plant the first day. Alate B. brassicae were more active than M. persicae : both species showed alternating periods of activity and quiescence.
At light intensities between 100 and 1000 f.c. there was little difference in flight frequency, but below 100 f.c. activity declined rapidly and apparently ceased with darkness.
Changes in relative humidity temporarily affected flight frequency, a change to a higher humidity retarding, a change to a lower increasing it. After adjusting to the change aphids flew readily at all humidities tested between 50 and 100% with temperatures below 80° F. (26.7°C.). The combination of high humidity and high temperature (90°F. = 32·2°C.) sometimes inhibited flight.
Changes of pressure often increased activity temporarily and flight frequency was greater under fluctuating pressure than under constant pressure.
It is concluded that changes in microclimate in crops are adequate to influence frequency of flight of aphids and consequently the spread of virus diseases.     

Toxicity of [14C] phorate applied systemically to the aphids Myzus persicae (Sulz) and Brevicoryne brassicae (l.) grown on brussels sprout plants

The effect of foliar application of [14C] phorate on the larvae of M. persicae and B. brassicae was studied. Regression analysis of the toxic components in the foliage and the probit kill showed that B. brassicae was more susceptible to the insecticide than M. persicae. The level of radioactivity extracted from the leaves, aphids and their honeydew shows that there was no correlation between the applied dose and the mortality. The amounts of honeydew produced by colonies before and after application of the insecticide were measured. The ratio of honeydew calculated shows that in M. persicae there was a slight increase in the honeydew production as the insecticide was increased from sublethal doses. At higher doses over the lethal range of the insecticide, honeydew production declined gradually in both the species.     

Differences in susceptibility to attack by Brevicoryne brassicae (L.) on brussels sprouts

The attack by alate and other forms of Brevicoryne brassicae (L.) and the amount of leaf malformation they caused on each of eighteen varieties or plant breeders' lines of brussels sprout were assessed in both 1967 and 1968 over the first 6 weeks following planting out. The results of the two years agreed closely despite differences in the dates of planting relative to aphid immigration. As a group, the Dutch varieties Roodnerf, Gravendeel, Tribune, Hercules and Castricum Glory were preferred by the incoming alates but suffered less damage than the least preferred inbred line, 3BL, of Cambridge No. 3. Some evidence of antibiosis was obtained, especially from a later experiment with three Cambridge inbred lines. However, in 1969 when there was a high rate of aphid multiplication on six varieties representing sprout types from Holland, Cambridge, Bedford and Ireland, any expression of host non-preference or antibiosis was soon obscured by the development of large aphid infestations and much damage. Under these circumstances, considerable varietal differences in tolerance to attack were found. Irish Elegance sustained most damage yet its most heavily attacked plants supported little more than half of the aphid numbers of equivalent plants of the most tolerant variety Darkmar 21. The Dutch varieties Vremo Inter and Seven Hills were almost as tolerant as Darkmar 21 and recovered quickly from damage. Darkmar 21 also showed high tolerance to attack in 1970, as did the Dutch varieties Rollo, Roodnerf and Stiekema, when large aphid infestations developed on the eight varieties then grown. Lack of plant tolerance to damage appeared to be associated with incurling foliage, a characteristic of Irish Elegance, Cambridge No. 1 and No. 3.     

Bottom‐up effects mediated by an organic soil amendment on the cabbage aphid pests Myzus persicae and Brevicoryne brassicae

Earthworm‐produced compost or vermicompost has been shown to increase resistance of plants to a variety of insect pests, but it is still unclear whether this resistance is dose dependent and whether the mechanisms responsible are the same for insect species with differing feeding habits and preferences. Therefore, we tested the effects of plants grown in various vermicompost concentrations (0, 20, 40, and 60%) on the preference and performance of generalist, Myzus persicae L., and specialist, Brevicoryne brassicae L. (both Hemiptera: Aphididae), aphid pests. Preference was evaluated with leaf disk (apterous) and whole plant (alate) choice assays. After 24 h of feeding, there was no significant negative effect on the feeding preference noted for apterae of either species of any of the treatments tested. To the contrary, apterae B. brassicae showed a significant preference for vermicompost treatments over control leaf disks. Alate M. persicae preferred alighting on control plants over vermicompost‐grown plants, but B. brassicae showed no preference toward any of the treatments tested. Both aphid species deposited significantly more nymphs on control plants than on those grown in 20% vermicompost. Furthermore, plants grown in soil amended with 20% vermicompost significantly suppressed mass accumulation, as well as numbers of adults and nymphs of both aphid species compared to controls. These data clearly show that vermicompost soil amendments can significantly influence pest aphid preference and performance on plants and that these effects are not dose dependent, but rather species and morph dependent.     

Effects of infection with the cabbage black ringspot strain of turnip mosaic virus on turnip as a host to Myzus persicae and Brevicoryne brassicae

When turnip plants with 3–7 leaves were inoculated with cabbage black ringspot virus (CBRSV) on the 3rd rough-leaf, symptoms only appeared on leaves that had been less than 15 mm long at the time of inoculation, although infection decreased the area and both fresh and dry weight of all leaves. Leaves were ‘aged’ by their appearance and placed in Leaf Age Categories (LACs). Leaves with symptoms senesced (‘aged’) prematurely. CBRSV-infection of cv. Green Top White did not change the distribution of populations of Myzus persicae between LACs, but increased the proportion of the plant suitable for colonisation. All suitable LACs were quickly colonised by adult apterae and nymphs. On CBRSV-infected plants the nymphal period was shorter, F₁ adults deposited larvae more frequently and the live body weight and tibial length of the F₂ generation was greater, than on healthy plants. The distribution of Brevicoryne brassicae populations on cv. Green Top White differed from that of M. persicae but was also unchanged by CBRSV-infection. On healthy plants the largest colonies were on mature leaves, so that on virus-infected plants premature senescence shortened the life of the colony. On CBRSV-infected plants the nymphal period was prolonged and the live weight of F₁ and F₂ adult apterae was less than on healthy plants. The differences between the biology of M. persicae and B. brassicae on CBRSV-infected cv. Green Top White were associated with the accelerated senescence of CBRSV-infected leaves. The possibility that CBRSV-infection might reduce the resistance of turnips to aphid infestation was tested. M. persicae and B. brassicae were cultured on two favourable and two less favourable cultivars. No improvement in population growth rate was found when the less favourable host cultivars were infected with CBRSV, but both aphid species weighed less and/or had smaller nymphal populations on cultivars showing the severest symptoms. These results are discussed in relation to the evolution of non-persistent virus transmission by aphids.     

The effect of Brevicoryne brassicae on leaf area, dry matter distribution and amino acids of the Brussels sprout plant

Twenty-six direct or derived criteria were assessed, eight wk after infestation, of the effect of an initial inoculum of 0, 5, 25 or 50 Brevicoryne brassicae on the growth and dry matter distribution of Brussels sprout plants. Twenty-four of these criteria showed significant (P > 0.05) effects of aphid infestation, though increasing aphid infestation above 5/plant had little further effect. Total fresh and dry weight (as well as of the components — leaves, stems and roots) was reduced by 25–730 with infestation, particularly so the root. The number of leaves as well as individual leaf area was reduced, and leaves on infested plants were less efficient assimilators. This could be explained by the drain on assimilates caused by the aphids, the leaf curling and the reduced density of stomata. The latter was also considered to account partly for the increased moisture content of infested plants. There was a general reduction in the concentration of leaf amino acids.     

The relative importance of glucosinolates and amino acids to the development of two aphid pests Brevicoryne brassicae and Myzus persicae on wild and cultivated brassica species

This paper describes an experiment to investigate the relative importance of glucosinolates and the availability of free amino acids in the phloem to the feeding behaviour and development of the specialist brassica aphid, Brevicoryne brassicae (L) (cabbage aphid) and the generalist, Myzus persicae, (Sultzer) (peach potato aphid). Aphid development was determined on brassica species and cultivated brassica varieties. Analysis of individual glucosinolates in the wild brassica species identified significant differences in their profiles and in their concentrations present in freeze dried leaves. Multiple regression analysis showed a significant correlation ( ) between the intrinsic rate of increase of B. brassicae and glucosinolate concentrations; four glucosinolates, 2-OH-3-butenyl, and 2-propenylglucosinolate, 3-methoxyindolyl and 4-pentenylglucosinolates accounted for 79% of the variation. The intrinsic rate of increase of M. persicae was less correlated, though still significant ( ); 3-indolyglucosinolate and 3-butenylglucosinolate accounted for 47% of the variation. Regression analysis also showed a correlation between phloem amino acid concentrations and the intrinsic rate of increase of B. brassicae ( ) but not of M. persicae. The concentration of the important amino acids tyrosine, alanine, leucine and glutamic acid, accounted for 43% of the variation in intrinsic rate of increase. Leaf water potential of the Brassica species showed no relationship with the intrinsic rate of increase of either B. brassicae or M. persicae.     

Shifts in host plant resistance to the cabbage aphid (Brevicoryne brassicae) exhibited by Brussels sprout plants

Seven cultivars of Brussels sprouts from different sources were tested for their relative degrees of resistance to the cabbage aphid (Brevicoryne brassicae) by estimating the Potential Increase Rate of aphid infestations. The cultivars Winter Harvest and Early Half Tall were selected as susceptible and resistant respectively and tested further. Glasshouse experiments indicated that the resistance of Winter Harvest to the aphids increased with age.
These relative levels of resistance were confirmed by population counts in the field. Aphid numbers were initially significantly higher on Winter Harvest than on Early Half Tall although significance was lost as the season progressed at a time when the characteristics responsible for the resistance mechanism ceased to be distinguishable between the two cultivars.
Analysis of the amino acid content of leaf tissues collected at monthly intervals through the aphid season enabled 'risk rating' predictions of aphid resistance to be made. These conformed with the shifts towards greater resistance as the plants aged noted in the glasshouse trials and endorsed the part played by amino acids in the mechanism of resistance. Total amino acid concentrations reached a peak during September and October then declined with generally lower levels of amino acids in Early Half Tall than Winter Harvest. This cultivar also possessed a greater concentration of those amino acids known to be favourable to B. brassicae although the difference between the two had diminished considerably by the end of September.
Analyses of the mustard oil glycoside levels in mature glasshouse-grown plants did not indicate a difference between the cultivars.     

番茄叶片光合作用对快速水分胁迫的响应

采用聚乙二醇(PEG-6000)溶液控制番茄根际水势和叶片离体的方式设置了水分胁迫处理,测算了光合诱导过程中净光合速率、暗呼吸速率和CO2补偿点等光合参数的变化.结果表明:在1000μmol·m-2·s-1光诱导下,水分胁迫处理的番茄叶片净光合速率(Pn)达到最大值所需时间缩短为对照的1/3,气孔导度(gs)快速增大为对照的1.5倍.水分胁迫处理的番茄叶片光饱和点(LSP)比对照降低了65％～85％,而光补偿点(LCP)比对照增加了75％ ～100％,缩小了番茄叶片利用光能的有效范围.水分胁迫处理的番茄叶片最大光合能力(Amax)低于对照40％以上,暗呼吸速率(Rd)增大了约45％.可见,快速水分胁迫处理使番茄叶片气孔迅速开放,光合诱导初始阶段消失.水分胁迫导致植物利用光能的效率和潜力降低是植物生产力下降的重要原因,而气孔调节是番茄适应快速水分胁迫的重要生理机制.