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1.
  • 1 The vine weevil Otiorhynchus sulcatus is a major pest of horticultural crops worldwide, with root‐feeding larvae causing most damage. Adult oviposition aboveground may therefore influence levels of damage as the larvae are relatively immobile after oviposition.
  • 2 The present study investigated feeding and oviposition behaviour on red raspberry Rubus idaeus using intact plants, ensuring that choices reflected the realistic differences in cultivar appearance and chemical composition. Previous studies investigating vine weevil feeding and oviposition on other crops have used excised plant material, which may inadvertently influence behaviour.
  • 3 Adult weevils significantly preferred to feed on particular cultivars in the choice experiment (e.g. Tulameen), although they consumed significantly more foliage (0.22–1.03 cm2/day) on different raspberry cultivars (e.g. Glen Moy, Glen Rosa and a wild accession) in no‐choice situations.
  • 4 In choice experiments, weevils tended to avoid laying eggs on some cultivars (e.g. Glen Moy and the wild accession). The number of eggs laid (1.91–4.32 eggs per day) did not, however, differ significantly between the cultivars in a no‐choice situation. Foliar nitrogen and magnesium concentrations were positively, although weakly, correlated with the total number of eggs laid.
  • 5 The present study highlights the importance of considering both choice and no‐choice tests when assessing crop susceptibility to attack because weevils may avoid feeding on certain cultivars (e.g. Glen Moy) when given a choice, although this would cause significant damage to such cultivars if they were grown in monoculture (i.e. when there is no alternative).
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2.
1. A substantial amount of research on host‐plant selection by insect herbivores is focused around the preference–performance hypothesis (PPH). To date, the majority of studies have primarily considered insects with aboveground life cycles, overlooking insect herbivores that have both aboveground and belowground life stages, for which the PPH could be equally applicable. 2. This study investigated the factors influencing the performance of the root‐feeding vine weevil (Otiorhynchus sulcatus) larvae and whether this was linked to the oviposition behaviour of the maternal adult living aboveground. 3. Maternal insects feeding aboveground reduced root biomass by 34% and increased root carbon by 4%. Larvae feeding on plants subjected to aboveground herbivory had reduced mass. Irrespective of the presence of maternal herbivory, larval mass was positively correlated with root biomass. 4. Larval mass was also reduced by conspecific larvae, previously feeding on roots (19% reduction). However, the mechanism underpinning this effect remains unclear, as in contrast to maternal herbivory aboveground, prior larval feeding did not significantly affect root biomass or root carbon concentrations. 5. Maternal insects did not distinguish between plants infested with larvae and those that were free of larvae, in terms of their egg‐laying behaviour. Conversely, maternal insects tended to lay eggs on plants with smaller root systems, a behaviour that is likely to negatively affect offspring performance. 6. The PPH is not supported by our findings for the polyphagous vine weevil feeding on the host plant raspberry (Rubus idaeus), and in fact our results suggest that there is the potential for strong parent–offspring conflict in this system.  相似文献   

3.
Black vine weevils, Otiorhynchus sulcatus (Fabricius) (Coleoptera: Curculionidae), are globally‐distributed polyphagous pests of many horticultural crops. We investigated how adult weevils were affected by host switching and, in particular, how host plant species nutritional and defensive chemistry affected subsequent host plant species selection and oviposition. Adults were fed one of three host plant species, blackcurrant [Ribes nigrum L. (Grossulariaceae)], raspberry [Rubus idaeus L. (Rosaceae)], or strawberry [Fragaria x ananassa Duchesne (Rosaceae)], throughout their pre‐reproductive periods and then subjected to behavioral choice assays with these plants. Foliar chemistry differed significantly among the three host plant species. Compared to raspberry and strawberry foliage, blackcurrant foliage was 13% lower in nitrogen, 3% higher in carbon, and 28% higher in phenolic compounds. Initial host plant species had a significant effect on weevil mortality, with more weevils dying when previously fed blackcurrant (12%) than strawberry (3%) or raspberry (0%) regardless of subsequent host. Initial host plant species also affected oviposition, with weevils laying only ca. two eggs per week when previously fed blackcurrant, compared to those on raspberry or strawberry (ca. 11 and 15 eggs per week, respectively). When given a choice, weevils discriminated among host plant species and tended to oviposit on plants on which they had previously fed, even when the plant was nutritionally inferior for egg production and adult survival. In contrast, feeding behavior was only affected by the current host plant species. Feeding and oviposition were related to leaf chemistry only in blackcurrant, as leaf consumption was negatively correlated with foliar carbon and zinc concentrations, and positively correlated with foliar phosphorus and potassium concentrations.  相似文献   

4.
5.
Detrimental effects of vines on tree growth in successional environments have been frequently reported. Little is known, however, about the relative importance of below and aboveground competition from vines on tree growth. The objective of this study was to quantify and compare the growth responses of Liquidambar styraciflua saplings to below and/or aboveground competition with the exotic evergreen vine, Lonicera japonica (Japanese honeysuckle), and the native deciduous vine, Parthenocissus quinquefolia (Virginia creeper). Soil trenching and/or vine-trellising were used to control the type of vine competition experienced by trees. Comparisons among untrenched treatments tested for effects of belowground competition. Comparisons among trenched treatments tested for effects of aboveground competition. After two growing seasons, Lonicera japonica had a greater effect on the growth of L. styraciflua than did P. quinquefolia. This effect was largely due to root competition, as canopy competition only had a negative effect on tree growth when it occurred in combination with root competition. Leaf expansion was consistently and similarly affected by all treatments which involved belowground competition.  相似文献   

6.
Five strawberry (Fragaria sp.) and five raspberry (Rubus ideaus L.) cultivars were evaluated for resistance to two spotted spider mite (Tetranychus urticae Koch.). Two methods of assessing the development of two spotted mite populations using detached leaves were compared. The number of eggs laid and mites which developed were compared. The strawberry cvs Hapil and Pegasus had significantly greater development of two spotted mite populations than the cvs Rhapsody, Symphony and Elsanta. The raspberry cv. Joan Squires had higher populations of two spotted mite whilst the raspberry cv. Leo the least, when compared with cvs Glen Clova, Glen Moy and Glen Prosen. Differences were observed in oviposition sites and mite distribution when comparing raspberries with strawberries. The method of assessing the populations development of two spotted mite which involved maintaining the cut leaf stem in water may be of potential use for studying population dynamics of both two spotted mite and possible predators over extended periods of time.  相似文献   

7.
Breeding in raspberry is time-consuming due to the highly heterozygous nature of this perennial fruit crop, coupled with relatively long periods of juvenility. The speed and precision of raspberry breeding can be improved by genetic linkage maps, thus facilitating the development of diagnostic markers for polygenic traits and the identification of genes controlling complex phenotypes. A genetic linkage map (789 cM) of the red raspberry Rubus idaeus has been constructed from a cross between two phenotypically different cultivars; the recent European cultivar Glen Moy and the older North American cultivar Latham. SSR markers were developed from both genomic and cDNA libraries from Glen Moy. These SSRs, together with AFLP markers, were utilised to create a linkage map. In order to test the utility of the genetic linkage map for QTL analysis, morphological data based on easily scoreable phenotypic traits were collected. The segregation of cane spininess, and the root sucker traits of density and spread from the mother plant, was quantified in two different environments. These traits were analysed for significant linkages to mapped markers using MapQTL and were found to be located on linkage group 2 for spines and group 8 for density and diameter. The availability of co-dominant markers allowed heterozygosities to be calculated for both cultivars.  相似文献   

8.
Six blackberry or hybrid berry cultivars and 19 raspberry cultivars were assessed for their infectibility with, and sensitivity to, graft inoculation with 10 distinct viruses found infecting Rubus in the UK. Cultivars were grafted with each of, two isolates of the pollen borne raspberry bushy dwarf virus (RBDV), five aphid borne viruses: black raspberry necrosis, raspberry leaf mottle (RLMV), raspberry leaf spot (RLSV), rubus yellow net and raspberry vein chlorosis (RVCV); and isolates of the nematode transmitted nepoviruses, arabis mosaic, raspberry ringspot, strawberry latent ringspot and tomato black ring. All tested cultivars were infectible with a resistance breaking isolate of RBDV but only about half of that number with the Scottish type isolate of the virus. The raspberry cvs Autumn Bliss, and occasionally Glen Garry and Glen Prosen, developed leaf yellowing symptoms following infection with RBDV, but none of the other infected cultivars showed obvious leaf symptoms when kept in a heated glasshouse during the growing season. All tested cultivars were infectible with each of the four viruses transmitted in nature by the aphid, Amphorophora idaei. Most were infected symptomlessly, but seven cultivars developed severe leaf spotting symptoms due to infection with RLMV or RLSV. All but one of the raspberry cultivars were infectible with RVCV, which is transmitted in nature by the aphid Aphis idaei, and almost all infected plants developed leaf symptoms; only one of the hybrid berry or blackberry cultivars tested was infected with RVCV. In tests with the four nepoviruses, all tested cultivars, except Tummelberry, were infectible with at least one or more of these viruses. However, cultivars responded differently to challenge inoculation with different isolates of individual nepoviruses. Several cultivars developed chlorotic leaf mottling following infection with some nepovirus isolates. The implications of these results for virus control are discussed in the light of the changing pattern of virus and virus vector incidence in the UK.  相似文献   

9.
Belowground communities can affect interactions between plants and aboveground insect communities. Such belowground–aboveground interactions are known to depend on the composition of belowground communities, as well as on the plant species that mediates these interactions. However, it is largely unknown whether the effect of belowground communities on aboveground plant–insect interactions also depends on genotypic variation within the plant species that mediates the interaction. To assess whether the outcome of belowground–aboveground interactions can be affected by plant genotype, we selected two white cabbage cultivars [Brassica oleracea L. var. capitata (Brassicaceae)]. From previous studies, it is known that these cultivars differ in their chemistry and belowground and aboveground multitrophic interactions. Belowground, we inoculated soils of the cultivars with either nematodes or microorganisms and included a sterilized soil as a control treatment. Aboveground, we quantified aphid [Brevicoryne brassicae (L.) (Hemiptera: Aphididae)] population development and parasitoid [Diaeretiella rapae (McIntosh) (Hymenoptera: Braconidae)] fitness parameters. The cultivar that sustained highest aphid numbers also had the best parasitoid performance. Soil treatment affected aphid population sizes: microorganisms increased aphid population growth. Soil treatments did not affect parasitoid performance. Cultivars differed in their amino acid concentration, leaf relative growth rate, and root, shoot, and phloem glucosinolate composition but showed similar responses of these traits to soil treatments. Consistent with this observation, no interactions were found between cultivar and soil treatment for aphid population growth or parasitoid performance. Overall, the aboveground community was more affected by cultivar, which was associated with glucosinolate profiles, than by soil community.  相似文献   

10.
The effect of photoperiod on nutritional quality and chemical composition of ‘Glen Ample’ red raspberry has been studied under controlled environment conditions. Long day (LD) conditions during fruit growth and maturation significantly increased fruit concentrations of ascorbic acid, total phenolics, organic acids such as malic, quinic and ellagic acids, as well as the ferric reducing activity power, while reducing the concentrations of soluble solids, sucrose and the sugar/acid ratio. The levels of total anthocyanin and oxalic acid were not significantly affected by photoperiod whereas semi‐quantitative analysis indicated a significant increase in the levels of some ellagitannins. The same responses were found whether LD was established by day‐length extension with low‐intensity light or by night interruption, thus demonstrating that the observed changes in nutritional quality of raspberry fruit was mediated through specific photoperiodic mechanism(s) independently of the daily light energy supply.  相似文献   

11.
Predicting how insect crop pests will respond to global climate change is an important part of increasing crop production for future food security, and will increasingly rely on empirically based evidence. The effects of atmospheric composition, especially elevated carbon dioxide (eCO2), on insect herbivores have been well studied, but this research has focussed almost exclusively on aboveground insects. However, responses of root‐feeding insects to eCO2 are unlikely to mirror these trends because of fundamental differences between aboveground and belowground habitats. Moreover, changes in secondary metabolites and defensive responses to insect attack under eCO2 conditions are largely unexplored for root herbivore interactions. This study investigated how eCO2 (700 μmol mol?1) affected a root‐feeding herbivore via changes to plant growth and concentrations of carbon (C), nitrogen (N) and phenolics. This study used the root‐feeding vine weevil, Otiorhynchus sulcatus and the perennial crop, Ribes nigrum. Weevil populations decreased by 33% and body mass decreased by 23% (from 7.2 to 5.4 mg) in eCO2. Root biomass decreased by 16% in eCO2, which was strongly correlated with weevil performance. While root N concentrations fell by 8%, there were no significant effects of eCO2 on root C and N concentrations. Weevils caused a sink in plants, resulting in 8–12% decreases in leaf C concentration following herbivory. There was an interactive effect of CO2 and root herbivory on root phenolic concentrations, whereby weevils induced an increase at ambient CO2, suggestive of defensive response, but caused a decrease under eCO2. Contrary to predictions, there was a positive relationship between root phenolics and weevil performance. We conclude that impaired root‐growth underpinned the negative effects of eCO2 on vine weevils and speculate that the plant's failure to mount a defensive response at eCO2 may have intensified these negative effects.  相似文献   

12.
Systemic activity of the neonicotinoids clothianidin, dinotefuran, and thiamethoxam and the anthranilic diamide chlorantraniliprole was tested against adult black vine weevils, Otiorhynchus sulcatus (F.) (Coleoptera: Curculionidae), on Astilbe, Euonymus, Heuchera, Rhododendron, Sedum, and Taxus. Insecticide treatments were applied to the soilless substrate of containerized plants. Bioassays were conducted 12 or 13, 26, and 42 d after treatment (DAT) and ran for 7 d; and feeding, mortality, and weight gain or loss by weevils were evaluated. Foliage was removed from test plants and then placed in arenas with adult black vine weevils. The neonicotinoids reduced feeding and weight gain by adult black vine weevils on most plant species with residual activity 42 DAT on some plant species. At 12 DAT, mortality was caused by the three neonicotinoids on Astilbe and by thiamethoxam on Sedum; and at 26 DAT dinotefuran caused mortality on Astilbe. Chlorantraniliprole reduced feeding on Taxus at 12 DAT, with no activity detected in other bioassays. Another set of bioassays was conducted to examine survival and fecundity of adult black vine weevils during prolonged feeding on Heuchera and Taxus systemically treated with dinotefuran or thiamethoxam. Bioassay procedures were similar to those described above, except they ran continuously for 56 d. Prolonged feeding on dinotefuran and thiamethoxam treated Heuchera and Taxus resulted in high mortality of adult black vine weevils and reduced fecundity. These studies show that the systemic activity of neonicotinoids is influenced by plant species and that systemic neonicotinoids have the potential to suppress black vine weevil populations in containerized nursery crops.  相似文献   

13.
Abstract. 1. In laboratory experiments, R.fagi made feeding holes in eighteen tree and shrub species, including beech, from seven plant families but few plants other than beech were eaten to any extent.
2. Overwintered adult weevils preferred beech to all these eighteen plants in spring.
3. Hawthorn and raspberry were the most acceptable plants if beech was not available at this time.
4. Field collected weevils from hawthorn had partly developed oocytes in early April, prior to beech bud burst.
5. New generation weevils sought alternative food sources in late June and July; raspberry was preferred to the youngest available beech leaves at this time.  相似文献   

14.
Abstract 1 The feeding preference of vine weevil, Otiorhynchus sulcatus (Fabricius), adults for foliage from 21 commercial cultivars of strawberries is investigated using binary choice tests with leaf disks, using ‘Honeoye’ as a standard against which all other cultivars are compared. ‘Delmarvel’, ‘Idea’, ‘Lester’, ‘Primetime’ and ‘Seneca’ are not preferred. 2 Variation in leaf nitrogen content is correlated with feeding preference, but does not explain all the variation, because outliers exist for both preferred and nonpreferred cultivars. 3 Removal of leaf hairs with adhesive tape permits their role in deterrence to be evaluated. Eight of the 21 cultivars have deterrent leaf hairs. Paradoxically, some highly preferred cultivars (e.g. ‘Latestar’ and ‘Tristar’) have deterrent leaf hairs, and four of the five nonpreferred cultivars lack significantly deterrent leaf hairs. 4 The 21 tested cultivars do not differ in their suitability for vine weevil larval development. 5 Differences in cultivar susceptibility to vine weevil in the field may involve interactions between the palatability and nutritional value of foliage, which influence fecundity and egg placement, and the tolerance of strawberry plants to larval feeding, determined by root biomass. The nonpreference traits found in the foliage of commercial cultivars of strawberry are present by chance. Accordingly, further selective breeding to enhance strawberry varietal tolerance to vine weevils may be possible.  相似文献   

15.
Cosme M  Stout MJ  Wurst S 《Mycorrhiza》2011,21(7):651-658
Root-feeding insects are important drivers in ecosystems, and links between aboveground oviposition preference and belowground larval performance have been suggested. The root-colonizing arbuscular mycorrhizal fungi (AMF) play a central role in plant nutrition and are known to change host quality for root-feeding insects. However, it is not known if and how AMF affect the aboveground oviposition of insects whose offspring feed on roots. According to the preference–performance hypothesis, insect herbivores oviposit on plants that will maximize offspring performance. In a greenhouse experiment with rice (Oryza sativa), we investigated the effects of AMF (Glomus intraradices) on aboveground oviposition of rice water weevil (Lissorhoptrus oryzophilus), the larvae of which feed belowground on the roots. Oviposition (i.e., the numbers of eggs laid by weevil females in leaf sheaths) was enhanced when the plants were colonized by AMF. However, the leaf area consumed by adult weevils was not affected. Although AMF reduced plant biomass, it increased nitrogen (N) and phosphorus concentrations in leaves and N in roots. The results suggest that rice water weevil females are able to discriminate plants for oviposition depending on their mycorrhizal status. The discrimination is probably related to AMF-mediated changes in plant quality, i.e., the females choose to oviposit more on plants with higher nutrient concentrations to potentially optimize offspring performance. AMF-mediated change in plant host choice for chewing insect oviposition is a novel aspect of below- and aboveground interactions.  相似文献   

16.
The importance of species richness to ecosystem functioning and services is a central tenet of biological conservation. However, most of our theory and mechanistic understanding is based on diversity found aboveground. Our study sought to better understand the relationship between diversity and belowground function by studying root biomass across a plant diversity gradient. We collected soil cores from 91 plots with between 1 and 12 aboveground tree species in three natural secondary forests to measure fine root (≤ 2 mm in diameter) biomass. Molecular methods were used to identify the tree species of fine roots and to estimate fine root biomass for each species. This study tested whether the spatial root partitioning (species differ by belowground territory) and symmetric growth (the capacity to colonize nutrient-rich hotspots) underpin the relationship between aboveground species richness and fine root biomass. All species preferred to grow in nutrient-rich areas and symmetric growth could explain the positive relationship between aboveground species richness and fine root biomass. However, symmetric growth only appeared in the nutrient-rich upper soil layer (0–10 cm). Structural equation modelling indicated that aboveground species richness and stand density significantly affected fine root biomass. Specifically, fine root biomass depended on the interaction between aboveground species richness and stand density, with fine root biomass increasing with species richness at lower stand density, but not at higher stand density. Overall, evidence for spatial (i.e. vertical) root partitioning was inconsistent; assumingly any roots growing into deeper unexplored soil layers were not sufficient contributors to the positive diversity–function relationship. Alternatively, density-dependent biotic interactions affecting tree recruitment are an important driver affecting productivity in diverse subtropical forests but the usual root distribution patterns in line with the spatial root partitioning hypothesis are unrealistic in contexts where soil nutrients are heterogeneously distributed.  相似文献   

17.
Plants are often simultaneously or sequentially attacked by multiple herbivores and changes in host plants induced by one herbivore can influence the performance of other herbivores. We examined how sequential feeding on the plant Plantago lanceolata by the aboveground herbivore Spodoptera exigua and the belowground herbivore Agriotes lineatus influences plant defense and the performance of both insects. Belowground herbivory caused a reduction in the food consumption by the aboveground herbivore independent of whether it was initiated before, at the same time, or after that of the aboveground herbivore. By contrast, aboveground herbivory did not significantly affect belowground herbivore performance, but significantly reduced the performance of later arriving aboveground conspecifics. Interestingly, belowground herbivores negated negative effects of aboveground herbivores on consumption efficiency of their later arriving conspecifics, but only if the belowground herbivores were introduced simultaneously with the early arriving aboveground herbivores. Aboveground–belowground interactions could only partly be explained by induced changes in an important class of defense compounds, iridoid glycosides (IGs). Belowground herbivory caused a reduction in IGs in roots without affecting shoot levels, while aboveground herbivory increased IG levels in roots in the short term (4 days) but only in the shoots in the longer term (17 days). We conclude that the sequence of aboveground and belowground herbivory is important in interactions between aboveground and belowground herbivores and that knowledge on the timing of exposure is essential to predict outcomes of aboveground–belowground interactions.  相似文献   

18.
The introduction into commerce of raspberry cultivars with major gene resistance to the large raspberry aphid, Amphorophora idaei, an important pest and virus vector on red raspberry in Europe, has been very effective both in decreasing pest numbers and greatly restricting infection with the viruses it transmits. However, biotypes of the aphid able to overcome these genes have developed in the field in recent years. Additionally, in field and laboratory tests, the response to aphid biotypes and recognised aphid strains of certain raspberry cultivars, such as Glen Prosen and Delight, differ markedly despite the fact that they are reputed to contain the same A. idaei‐resistance gene, A1. In attempts to understand the reasons for this difference in response, analysis was made of the segregation of progeny seedlings from crosses between A. idaei‐resistant and ‐susceptible cultivars to two recognised strains of the aphid. These studies showed that, as expected, cv. Autumn Bliss contained the A. idaei‐resistance gene, A10, and cvs Delight and Glen Prosen each contained the A. idaei‐resistance gene, A1. When progeny seedlings were assayed in a heated glasshouse as young plants and in an unheated Tygan house as 1 m tall plants, the segregation ratios for resistance and susceptibility to A. idaei were largely unchanged. However, when the resistance of individual progeny plants was assessed, c. 37% of the putative gene A1‐containing progeny and 9–23% of the putative gene A10‐containing progeny, behaved differently in these two environments. Experiments involving an A. idaei‐resistant and ‐susceptible parent cultivar showed that shading plants increased their susceptibility to A. idaei colonisation. Whilst this shading effect has implications for experimentally detecting A. idaei‐resistant progeny in segregating raspberry seedlings, it does not explain the difference in field resistance to A. idaei of cvs Delight and Glen Prosen. Such differences in the field seem best explained by the presence in these cultivars of ‘minor’ genes for A. idaei resistance and/or susceptibility that influences the effectiveness of gene A1.  相似文献   

19.
松嫩平原不同生育期虎尾草无性系构件生长与生物量分配   总被引:1,自引:0,他引:1  
以松嫩平原不同生育期的虎尾草无性系为研究对象,分别在虎尾草无性系拔节期和完熟期进行大样本取样,并对其地上、地下各构件的数量性状及生物量分配进行统计分析,研究虎尾草在不同生育期的生长特性及生长策略.结果表明:虎尾草株高、总根长、总根表面积、根体积、地上生物量、地下生物量、总生物量在两个生育期之间均存在显著差异.在两个生育期,地上、地下生物量分配与总生物量呈显著幂函数异速生长关系.在拔节期,总生物量与分株数、总根长、总根表面积和根体积呈显著线性同速生长关系;而在完熟期,均呈显著幂函数异速生长关系.虎尾草无性系在不同生育期存在着不同的生长策略,在拔节期主要采取的是无性系外部空间的优先扩展策略,而在完熟期主要采取的是无性系内、外部空间兼顾的补充和扩展策略.  相似文献   

20.
Ripening-related changes in raspberry cell wall composition and structure   总被引:1,自引:0,他引:1  
Cell walls were prepared from the fruit of two cultivars of raspberry at three stages of ripening; green, white and red (ripe). The cultivars. Glen Clova and Glen Prosen, are subjectively classified, at harvest by growers, as soft and firm fruit, respectively. The cell walls were analysed for neutral sugar composition, uronic acid content, degree of methyl esterification, lignin and ferulic acid-derived dehydrodimers. Solid-state 31C NMR and diffuse reflectance infrared (DRIFT) spectra were acquired for the cell wall residues. For both cultivars the progression from green to white produced minimal changes, save for a reduction in pectin. NMR analyses indicated that the solubilized pectin was acetylated. Progression to the red (ripe) stage, in both cultivars, was accompanied by a reduction in the ordered cellulose and a dramatic reduction in pectin content and the degree of methyl-esterification. Significantly, the softer fruit (Glen Clova) exhibited greater reductions in both parameters, implicating increased pectin hydrolysis, as one of the main factors contributing to the difference in firmness between the cultivars. A relative increase in cell wall-associated protein was seen at the red stage. The nature and function of the protein(s) are, as yet unknown.  相似文献   

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