Evaluation of induced responses, insect population growth, and host-plant fitness may change the outcome of tests of the preference-performance hypothesis: a case study

The preference‐performance hypothesis predicts that insect preference should correspond to host suitability for offspring development. We studied the pattern of within‐plant preference in the aphid Sipha flava and its consequences for offspring performance on the host‐plant Sorghum halepense, regarding the role of induced responses of plants to aphid feeding. The consequences of within‐plant preference on aphid population growth and host‐plant traits were also evaluated. Our results showed that winged and wingless aphids preferred to settle on mature rather than young leaves. In contrast, aphid individual growth rate was higher on young leaves when compared with mature leaves, suggesting that the outcome of this test rejected the preference‐performance hypothesis. However, the inclusion of the factor ‘previous aphid infestation’ changed the outcome from a maladaptive choice to a neutral one. Thus, individual growth rates of S. flava increased when aphids developed on leaves that had been previously infested. Interestingly, aphid growth rate on previously infested leaves did not differ between young and mature leaves. On the other hand, aphid population reproductive rate was higher and the percentage of winged aphids lower when infestation occurred on mature rather than young leaves. Aphid infestation reduced plant and shoot biomass, and increased leaf mortality. These negative effects on plant traits related to plant fitness were greater when aphid infestation occurred on young leaves. Likewise, whereas infestation on mature leaves did not cause a significant reduction in the number of flowering plants compared with control plants, aphid infestation on young leaves did reduce the number of plants at the flowering stage. Consequently, if both the reproductive rate of aphids in the mid‐term, and host‐plant fitness are taken into account, the results indicate that aphid preference for mature leaves may be an adaptive choice, thus supporting the preference‐performance hypothesis.     

The influence of black bean aphid, Aphis fabae Scop., and its honeydew on leaf growth and dry matter production of sugar beet

A. fabae populations, started at the 3–4 leaf-stage of sugar beet in the glasshouse and peaking at 3000 individuals per plant, reduced leaf area by 64% at the 14 leaf-stage. The size of the heavily-infested leaves number 5 to 10 was reduced by 80% or more. The rate of leaf growth regained normal values after the aphid populations collapsed, but the infested plants did not make up for the decrease in leaf area production that had been incurred during the infestation. Total dry matter production over a period of 15 weeks was reduced by 47%. Honeydew had no effect on leaf size or dry matter production. Sugar beet plants in the field became infested with A. fabae at the 2–3, 4–5 and 6–8 leaf stages. Maximum populations of 800, 2100 and 2200 aphids per plant were recorded, respectively. The pertinent reductions in leaf area were 91%, 67% and 34% at the 10–12 leaf-stage and 79%, 65% and 14% at harvest while the total dry matter produced was reduced by 91%, 79% and 16%. Neighbouring plants of the early-infested sugar beet plants gained significantly higher weights than control plants. Honeydew had no effect on leaf area or dry matter production. The consequences of these results for our understanding of Aphis fabae injury in sugar beet and aphid control in the field are discussed.     

The effect of plant age and infection with virus yellows on the survival of Myzus persicae on sugar beet

Survival of Myzus persicae confined in clip-cages on mature leaves of sugar beet declined as the plants aged. Death of aphids was often preceded by the appearance of a black deposit in the aphids' stomachs, which may have been the cause of death. Both the rate of death and the proportion of aphids dying with black deposits was significantly less when plants were infected with beet yellows virus or beet mild yellowing virus, by comparison with healthy plants. The implication of these phenomena on the onset of mature plant resistance is discussed.     

Analyses of the stomach deposit that develops in Myzus persicae feeding on sugar beet

Abstract .The white deposit, commonly found in the stomach of aphids that feed on sugar beet ( Beta vulgaris ssp. saccharifera L), turns a dark colour just prior to the death of the aphid, suggesting that chemical changes causing the deposit to darken may be associated with the death of the aphid. Chemical analyses of the white and dark stomach deposits by HPLC after hydrolysis, elemental analysis and FAB-MS, showed that the dark deposit is formed after the loss of hydrogen and oxygen from the organic white deposit, and that it appears to be a complex that is not dominated by any given compound. These results contrast with those of previous studies, in which the white precipitate was described as a polysaccharide or a mucopolysaccharide.     

The influence of black bean aphid, Aphis fabae Scop., and its honeydew on the photosynthesis of sugar beet

The effect of black bean aphids on the photosynthesis of sugar beet plants was studied under glasshouse and field conditions. The presence of up to several hundred aphids per leaf had no significant effect on CO₂ exchange rates over a range of light intensities between complete darkness and light saturation. Artificially prepared honeydew, sprayed onto leaves in the same amounts and composition as was found on severely aphid-infested plants, covered 30% of the stomata on the upper epidermis but did not significantly alter the rate of photosynthesis of these leaves in the light or the rate of respiration in the dark. The stomata on the lower epidermis were uncovered and functional. High pressure liquid chromatography of aphid-produced honeydew detected 20 different amino-acids. Three amino-acids, aspartic acid, glutamic acid and gluta-mine, made up the bulk of the amino-acid weight in the honeydew produced on young plants, up till the 8 leaf-stage. In the 10 to 12 leaf-stage, several different amino-acids occurred in substantial amounts. The amino-acids to sugars ratio of the honeydew produced by the aphids decreased strongly as the sugar beet plants aged: from 1:6 in plants with 3 or 4 leaves to 1:25 in plants having 10 to 12 leaves.     

Influence of long-term drought stress on osmolyte accumulation in sugar beet (Beta vulgaris L.) plants

Accumulation of various osmolytes was examined in plants of sugar beet cv. Janus grown under two soil water treatments: control (60% of the field water capacity; FWC) and drought (30–35% FWC). The water shortage started on the 61st day after emergence (DAE), at the stage of the beginning of tap-roots development and was imposed for 35 days. Osmotic potential of sugar beet plant organs, particularly tap-roots, was decreased significantly as a consequence of a long-term drought. Water shortage reduced univalent (K⁺, Na⁺) cations concentrations in the petioles and divalent (Ca²⁺, Mg²⁺) ions level in the mature and old leaves. Cation concentrations in the tap-roots were not affected by water shortage. The ratio of univalent to divalent cations was significantly increased in young leaves and petioles as a consequence of drought. Long-term water deficit caused a significant reduction of inorganic phosphorus (P_i) concentration in young and old leaves. Under the water stress condition, the concentration of proline was increased in all individual plant organs, except proline concentration in the youngest leaves. Drought treatment caused a significant increase of glycine betaine content in shoot without any change in tap-roots. Glucose concentrations were significantly increased only in tap-roots as the effect of drought. In response to water shortage the accumulation of sucrose was observed in all the examined leaves and tap-roots. Overall, a long-term drought activated an effective mechanism for osmotic adjustment both in the shoot and in the root tissues which may be critical to survival rather than to maintain plant growth but sugar beet organs accumulate different solutes as a response to water cessation.     

Iron deficiency causes changes in chlorophyll fluorescence due to the reduction in the dark of the Photosystem II acceptor side

Iron deficiency was found to affect the redox state of the Photosystem II acceptor side in dark-adapted, attached leaves of sugar beet (Beta vulgaris L.). Dark-adapted iron-deficient leaves exhibited relatively high F_o and F_pl levels in the Kautsky chlorophyll fluorescence induction curve when compared to the iron-sufficient controls. However, far-red illumination led to marked decreases in the apparent F_o and F_pl levels. Modulated fluorescence showed that far-red light decreased the fluorescence yield to the true F_o levels by increasing photochemical quenching, without inducing changes in the level of non-photochemical quenching. In dark-adapted, iron-deficient leaves, far-red illumination induced a faster fluorescence decay in the µs-ms time domain, indicating an improvement in the electron transport after the primary quinone acceptor in the reducing side of Photosystem II. All these data indicate that in iron-deficient leaves the plastoquinone pool was reduced in the dark. The extent of the plastoquinone reduction in sugar beet depended on the chlorophyll concentration of the leaf, on the time of preillumination and on the duration of dark adaptation. The dark reduction of plastoquinone was observed not only in sugar beet but also in other plant species affected by iron deficiency both in controlled conditions and in the field.     

Response of Solanum tuberosum to Myzus persicae infestation at different stages of foliage maturity

Young leaves of the potato Solanum tuberosum L. cultivar Kardal contain resistance factors to the green peach aphid Myzuspersicae （Sulzer） （Hemiptera： Aphididae） and normal probing behavior is impeded. However, M. persicae can survive and reproduce on mature and senescent leaves of the cv. Kardal plant without problems. We compared the settling ofM. persicae on young and old leaves and analyzed the impact of aphids settling on the plant in terms of gene expression. Settling, as measured by aphid numbers staying on young or old leaves, showed that after 21 h significantly fewer aphids were found on the young leaves. At earlier time points there were no difference between young and old leaves, suggesting that the young leaf resistance factors are not located at the surface level but deeper in the tissue. Gene expression was measured in plants at 96 h postinfestation, which is at a late stage in the interaction and in compatible interactions this is long enough for host plant acceptance to occur. In old leaves of cv. Kardal （compatible interaction）, M. persicae infestation elicited a higher number of differentially regulated genes than in young leaves. The plant response to aphid infestation included a larger number of genes induced than repressed, and the proportion of induced versus repressed genes was larger in young than in old leaves. Several genes changing expression seem to be involved in changing the metabolic state of the leaf from source to sink.     

The influence of size and duration of aphid infestation on host plant quality, and its effect on sugar beet yellowing virus epidemiology

This paper studies the influence of previous infestation on the host quality of sugar beet (Beta vulgaris L.) for aphids and the influence of previous infestation on sugar beet yellowing virus epidemiology. Sugar beet previously infested with Myzus persicae (Sulzer) or Aphis fabae Scopoli (Homoptera: Aphididae) had an improved host quality for subsequently infesting aphids of the same species. There was a significant negative relationship between the number of M. persicae infesting a plant and the proportion of those that died with a dark deposit in their stomachs, and a significant positive relationship between the number that settled on a plant and the number that infested it previously. Nymphs feeding on previously infested plants grew more rapidly than those on control plants. The beneficial effect of previous infestation persisted for at least 2 weeks and prolongation of the infestation beyond 2 weeks was of no further benefit to the aphids. Field grown sugar beet, previously colonised by M. persicae, was more susceptible to natural infestation by M. persicae up to 5 days after exposure. Previously infested plants were also more susceptible to infection with beet mild yellowing virus (BMYV) but not beet yellows virus (BYV), suggesting that the aphids on the previously infested sugar beet settled more readily and were more inclined to feed (and thus transmit BMYV) than aphids on the previously uninfested plants. The consequences for the control of sugar beet yellowing virus vectors are discussed.     

超表达AVP1基因提高甜菜的耐低磷和耐盐抗旱性

为探讨H⁺-焦磷酸酶编码基因对甜菜磷吸收和抗性的影响,实现优良基因在甜菜基因工程中的利用,研究在甜菜中超表达拟南芥液泡膜H⁺-焦磷酸酶编码基因AVP1,对转基因甜菜分析其耐低磷、耐盐性和抗旱性。结果显示,AVP1基因在甜菜植株的叶片和块根中表达,且在逆境胁迫下增强表达量响应胁迫;低磷处理条件下,转基因甜菜与野生型甜菜相比具有更高的含磷量,可提高甜菜对磷的吸收利用效率;干旱、盐胁迫处理条件下,AVP1基因在转基因甜菜中显著上升,在盐胁迫或干旱处理条件下,转基因植株的生长受抑程度相对较轻。随着盐和干旱胁迫的加剧,转基因植株体内MDA含量与野生型植株相比较低而脯氨酸含量显著增加,AVP1基因可通过减轻逆境对甜菜细胞膜的损伤及提高甜菜细胞的渗透调节能力,进而增强甜菜对高盐和干旱胁迫的抗性。     

Effect of surplus glucose on physiological and biochemical characteristics of sugar beet leaves in relation to the age of the leaf and the whole plant

Effect of surplus glucose on physiological and biochemical parameters of leaves of different age was investigated in sugar beet (Beta vulgaris L., subsp. saccharifera) plants in the stages of vegetative growth (SVG). Early and late SVG were differentiated by the ratio between the weights of roots and aboveground organs (0.10 and 0.35, respectively). The excess of Glu was produced by incubation of the disks excised from detached leaves in water or 0.1 M Glu at radiant flux density of 250 μmol/(m² s) with the light regime pattern described as night/day/night/light (8/16/8/3 h). In all the leaf disks incubated in water and glucose solution, the content of Glu and other soluble carbohydrates considerably increased as compared with their content in the leaves they were taken from. After disk incubation in water and glucose solution, the content of chlorophyll (a + b) rose as compared with its level in respective leaves in early SVG; in late SVG, it declined. In early SVG, the rate of the O₂ photosynthetic evolution (Ph) in the ageing leaves under saturating concentration of NaHCO₃ after incubation in water and Glu solution declined more considerably than in young leaves. In late SVG, incubation of leaf disks in water and Glu solution weakly affected P _n. The rate of O₂ dark consumption in the leaf disks of all the types of treatment increased after incubation in water and especially in Glu solution. Activity of soluble carbonic anhydrase (sCA) in the extracts from young leaves in early SVG after their incubation in water and Glu solution was essentially the same, but after the incubation of ageing leaves in Glu solution, it reliably decreased. In late SVG, sCA activity sharply decreased after incubation in water and Glu solution irrespective of the leaf age. In late SVG, activity of Rubisco in the young leaves did not change after their incubation in water but decreased after incubation of the leaves of the three ages in Glu solution. In early SVG, nonphotochemical fluorescence quenching (NPQ) in the young intact leaf was lower than in the ageing leaf, and after leaf incubation in water and Glu solution, it rose. In late SVG, the value of NPQ was greater than in early SVG and, in contrast to the leaves of early SVG, it declined after leaf incubation; in water, this decline was more pronounced than in the Glu solution. In early SVG, efficient quantum yield of photosystem II (PSII) was much greater than in late SVG and it declined in the leaves incubated with Glu. It was concluded that surplus Glu can maintain biosynthetic processes in the young leaves of young sugar beet plants (trophic function). A decline in the level of chlorophyll and the activities of sCA and Rubisco in the course of leaf development and senescence is considered as a symptom of the suppression of biosynthesis of proteins of chlorophyll-protein complexes and the enzymes (Rubisco and sCA).     

Reproduction of sugar beet mosaic and tobacco mosaic viruses in anthocyanized beet plants

During our studies on the interaction of anthocyanins and plant virus diseases, reproduction of sugar beet mosaic (SBMV) and tobacco mosaic viruses (TMV) was investigated. Experiments were carried out in leaves of sugar beet,Beta vulgaris cv. Dobrovicka N and its spontaneous anthocyanized mutant. SBMV induces a systemic infection while TMV is responsible for primary local symptoms in sugar beet leaves only. Our quantitative analyses onAmaranthus caudatus L. andChenopodium quinoa Wilid. showed a significant decrease in concentration of SBMV in juice extracted from anthocyanized beet plants as compared with extracts from normal green infected plants. Significant differences were also obtained when SBMV — containing juice was tested in mixtures with healthy extracts from anthocyanized and normal green plants. Also the intensity of TMV symptoms in beet leaves was considerably decreased in leaves of antho-eyanized plants.     

Some factors affecting the concentrations of sugars in leaves of inbred sugar-beet lines

In the glasshouse, large differences in concentrations of glucose and sucrose were observed between leaves of inbred sugar-beet lines that are known to differ from each other in resistance to pests and diseases. Differences between these lines in concentrations of fructose, glucose and sucrose were more pronounced in the petiole than in the lamina. The concentrations of glucose and fructose were lower in the first two leaves (primary leaves) than in those produced subsequently (secondary leaves); both types of leaf contained similar concentrations of sucrose. Secondary leaves from plants that had been kept in darkness for 17 h contained less fructose, glucose and sucrose than those of plants kept in sunlight for 5 h. Longer period of darkness lowered the concentrations of monosaccharides in the leaves further but did not affect the sucrose content. Primary and secondary leaves from the same sugar-beet plant often differ in non-race-specific resistance to pests and diseases; and darkness can affect suscetibility of beet to downy mildew and to the aphid Myzus persicae. The results of the sugar determinations therefore support the hypothesis that the concentrations of certain carbohydrates in sugar-beet leaves are important in non-race-specific resistance to pests and diseases.     

Development of Ramularia Leaf Spot on Sugar Beet as Influenced by Temperature and the Age of the Host Plant

A method of inoculating sugar beet plants (Beta vulgaris L.) with Ramularia beticola Faut. & Lamb, is described. Following inoculation, disease development in relation to temperature and plant age was studied for more than a month. The incubation period was 18 days at 10°C compared to 14 days at 17°C. At 25°C no symptoms appeared. Both temperature and plant age significantly influenced disease level and rate of disease development. Plants incubated at 17°C were more severely diseased 33 days after inoculation than plants incubated at 10°C. Young plants (3 weeks at inoculation) Were more susceptible than older plants (5 and 8 weeks at inoculatson) under growth chamber conditions. In the field, symptoms of Ramularia leaf spot appear relatively late in the season and young leaves are rarely attacked. The inconsistency of these observations is discussed.     

Characterization of the Xanthophyll Cycle and Other Photosynthetic Pigment Changes Induced by Iron Deficiency in Sugar Beet (Beta vulgaris L.)

In this work we characterize the changes induced by iron deficiency in the pigment composition of sugar beet (Beta vulgaris L.) leaves. When sugar beet plants were grown hydroponically under limited iron supply, neoxanthin and β-carotene decreased concomitantly with chlorophyll a, whereas lutein and the carotenoids within the xanthophyll cycle were less affected. Iron deficiency caused major increases in the lutein/chlorophyll a and xanthophyll cycle pigments/chlorophyll a molar ratios. Xanthophyll cycle carotenoids in Fe-deficient plants underwent epoxidations and de-epoxidations in response to ambient light conditions. In dark adapted Fe-deficient plants most of the xanthophyll cycle pigment pool was in the epoxidated form violaxanthin. We show, both by HPLC and by in vivo 505 nanometers absorbance changes, that in Fe deficient plants and in response to light, the de-epoxidated forms antheraxanthin and zeaxanthin were rapidly formed at the expense of violaxanthin. Several hours after returning to dark, the xanthophyll cycle was shifted again toward violaxanthin. The ratio of variable to maximum chlorophyll fluorescence from intact leaves was decreased by iron deficiency. However, in iron deficient leaves this ratio was little affected by light conditions which displace the xanthophyll cycle toward epoxidation or de-epoxidation. This suggests that the functioning of the xanthophyll cycle is not necessarily linked to protection against excess light input.     

Effects of sucrose sprays and darkness on aphid colonization of sugar beet and on aphid transmission of yellowing viruses

In the glasshouse, adult, apterous Myzus persicae (Sulz.) and Aphis fabae Scop, settled better and deposited more larvae on sucrose-sprayed sugar-beet plants than on water-sprayed plants. M. persicae settled badly and deposited few larvae on plants that were kept in the dark before or after infestation. The effects of darkness on aphids were reduced by spraying the host plants with 10% solutions of sucrose before infestation. Viruliferous M. persicae transmitted beet yellows virus (BYV) and beet mild yellowing virus (BMYV) less efficiently to dark-treated plants than to those grown in normal daylight. Spraying sugar beet with sucrose before inoculation with viruliferous M. persicae increased the proportion of successful BYV transmissions but only when the plants were dark-treated. The effects of sucrose and darkness on settling and larviposition of aphids and on virus transmission may be related to changes in the concentration of carbohydrates, particularly sugars, in the leaves.     

The effects of Beet mild yellowing virus and Beet chlorosis virus on the yield of UK field-grown sugar beet in 1997,1999 and 2000

The separate effects of the aphid‐transmitted poleroviruses; Beet mild yellowing virus (BMYV) and Beet chlorosis virus (BChV), on the yield of field‐grown sugar beet were studied following different inoculation dates from May to July in 1997,1999 and 2000. Each sugar beet plant within the appropriate plots was infected with virus using at least 10 wingless viruliferous Myzus persicae per plant. In all 3 years, overall yield losses caused by BMYV were negatively correlated with time of infection with early season (May) inoculations causing 18–27% losses in sugar yield but late season losses only 4–15%. BChV decreased the sugar yield and sugar content of beet following early season inoculations, although the effects on sugar yield were more variable (range 8–24%) and the virus appeared to be less damaging compared to BMYV. However, inoculations with BChV in July of each year caused greater root and sugar losses than inoculations with BMYV at that time. Both poleroviruses increased the sodium content of the roots early in the season, although neither virus had an effect on potassium levels at any stage.     

Early-season predation impacts the establishment of aphids and spread of beet yellows virus in sugar beet

The potential of predators to impact the establishment of aphid vectors and the spread of beet yellows virus in sugar beet was examined. Myzus persicae carrying beet yellows virus (BYV) were released on six interior sites and six edge sites in each of four fields at the end of May. Aphids established at low densities and BYV was spread in circular patches around the infested plants at all sites. The number of diseased plants per patch at the end of September ranged from a field-average of 130 to 210 in the four fields. There was a weak tendency towards better aphid establishment and greater virus spread in fields in less complex landscapes. Edge sites had less virus spread than interior sites in one field, more virus spread in two other fields, and there was no statistically significant difference in the fourth field. In the field where virus spread was lowest at edge sites, we used predator exclosure and direct observation to manipulate and quantify the effects of early season predation. On a warm day in early June, 81% ofAphis fabae exposed to predators on young beet plants disappeared during a 24 h period, compared to 10% of aphids protected by clipcages. Intermediate levels of predator exclusion, allowing aphids to walk away but restricting predator access, showed that predation was responsible for aphid disappearance.Cantharis lateralis L. (Coleoptera: Cantharidae) was the most frequently observed foliar predator (>90%). It was found eating aphids on several occasions. The incidence of predators was 1.8 per plant per h in the field interior and 3.8 per plant per h. near the edge. In the same field, aphids and virus were released in six edge and six interior sites, that were surrounded by 0.5 m high plastic open-top barriers (‘exclosures’). Pitfall trapping inside the barriers reduced potential soil predator densities to ca. one-tenth of the open field level and arrivals of flying predators were reduced. Inside the exclosures, aphid establishment was enhanced, and virus spread at exclosure sites was increased by about 50% compared to open sites. Foliar and pitfall sampling yielded the following predators:Cantharis lateralis, C. rufa L. (Coleoptera: Cantharidae),Coccinella septempunctata L.,C. undecimpunctata L. (Coleoptera: Coccinellidae),Pterostichus cupreus (L.),Harpalus rufipes (de Geer),Patrobus atrorufus (Strom),Trechus quadristriatus (Schrk.),Bembidion lampros (Herbst) (Coleoptera: Carabidae). In a laboratory no-choice trial (with 10M. persicae /day offered), each of these species ate aphids with consumption rates varying from 1.7 to 9.2 aphids/day. The results show that early predation substantially impacted aphid establishment in one field, and resulted in reduced virus spread. Results in the other fields show that these results cannot be easily generalized.     

Limitations for finding a linear relationship between chlorophyll content and photosynthetic activity

In experiments with fodder cabbage and sugar beet the influences were tested which restrict the finding of a uniform linear dependence between the chlorophyll content and photosynthetic rate. The main factor is most probably the irradiation during the growth of plants and the measuring of photosynthetic rate. Changes in the density and length of irradiation with the proceeding spring season are reflected in the decrease of unit chlorophyll activity in all leaves of the plant. Such differences were not observed in plants grown in winter months at a low greenhouse irradiation density. High irradiation density during the measurement of photosynthesis ensures the finding of high correlation of photosynthetic rate with the chlorophyll amount per unit area of leaf. With a low irradiation density, or in thick leaves, the lower layers of chloroplasts are not saturated with radiation, and hence a higher correlation coefficient is found by relating photosynthetic rate to the percentage of chlorophyll in dry weight than to the quantity of chlorophyll per unit area of leaf blade. In some experiments higher activity of chlorophylls was found in young leaves before they have reached the state of “photosynthetic maturity” than after this turning point. This difference diminished by relating the photosynthetic rate to chlorophylla content instead of the total amount of chlorophylls (a+b). High activity of chlorophylls in very young leaves and great individual variability within the experimental plant groups caused by uneven cultivation conditions are the basis of apparent sigmoid character of average curves for the chlorophyll-photosynthesis relation.