METABOLISM OF PLANT GROWTH REGULATORS

Comparison of the metabolism of ¹⁴C carboxyl-labelled 2,4-D in detached leaves of different varieties of apple showed that the 2,4-D-resistant Cox decarboxylated 57% of the 2,4-D in the leaf in 92 hr., whereas the susceptible Bramley's Seedling decarboxylated only 2% in the same time. Comparatively high rates of decarboxylation were found with all varieties of apple having Cox in their parentage; varieties unrelated to Cox showed low rates of decarboxylation. Similar high rates of decarboxylation were found in resistant varieties of strawberry and in Syringa vulgaris. Sixteen other species tested showed low rates of decarboxylation, irrespective of whether they were resistant or susceptible to 2,4-D.
The rate of decarboxylation of 2,4-D in the mature strawberry leaf was proportional to the amount of 2,4-D in the leaf. The rapid fall in decarboxylation rate in a detached leaf was due chiefly to reduction in the amount of 2,4-D present, and not to any falling off in the metabolic activity of the leaf. Young leaves decarboxylated 2,4-D at about half the rate of fully grown or senescent leaves. Oxidation of the methylene carbon proceeds at less than half the rate of that of the carboxyl carbon and is accompanied by an increase in the phenol content of the leaf which agrees closely with the amount of 2,4-D demethylated.
As previously reported for currants, part of the 2,4-D entering the leaf was bound and part inactivated by the formation of water-soluble compounds. The reaction of strawberry variety Talisman to 2,4,5-trichlorophenoxyacetic (2,4,5), 2-chloro-phenoxyacetic (2-CPA), 4-chlorophenoxyacetic (4-CPA) and 2-methyl-4-chlorophenoxyacetic (MCPA) acids was similar to that previously reported for red currant. Leaves of Cox's Orange Pippin apple and Talisman strawberry were able to decarboxylate all these phenoxy-acids except 2-CPA.     

Down-regulation of the strawberry Bet v 1-homologous allergen in concert with the flavonoid biosynthesis pathway in colorless strawberry mutant

Proteomic screening of strawberry (Fragaria ananassa) yielded a 58% success rate in protein identification in spite of the fact that no genomic sequence is available for this species. This was achieved by a combination of MALDI-MS/MS de novo sequencing of double-derivatized peptides and indel-tolerant searching against local protein databases built on both EST and full-length nucleotide sequences. The amino acid sequence of a strawberry allergen, homologous to the well-known major birch pollen allergen Bet v 1, was partially determined. This strawberry allergen, named Fra a 1 according to the nomenclature for allergen proteins, showed sequence identity of 54 and 77%, respectively, with corresponding allergens from birch and apple. Differential expression, as evaluated by 2-D DIGE, occurred in 10% of protein spots when red strawberries were compared to a colorless (white) strawberry mutant. White strawberries, known to be tolerated by individuals affected by allergy, were found to be virtually free from the strawberry allergen. Also several enzymes in the pathway for biosynthesis of flavonoids, to which the red color pelargonidin belongs, were down-regulated. This approach to assess differential protein expression without access to genomic sequence information can also be applied to other crop plants and phenotypic traits.     

Common protein biomarkers assessed by reverse phase protein arrays show considerable intratumoral heterogeneity in breast cancer tissues

Proteins are used as prognostic and predictive biomarkers in breast cancer. However, the variability of protein expression within the same tumor is not well studied. The aim of this study was to assess intratumoral heterogeneity in protein expression levels by reverse-phase-protein-arrays (RPPA) (i) within primary breast cancers and (ii) between axillary lymph node metastases from the same patient. Protein was extracted from 106 paraffin-embedded samples from 15 large (≥3 cm) primary invasive breast cancers, including different zones within the primary tumor (peripheral, intermediate, central) as well as 2-5 axillary lymph node metastases in 8 cases. Expression of 35 proteins including 15 phosphorylated proteins representing the HER2, EGFR, and uPA/PAI-1 signaling pathways was assessed using reverse-phase-protein-arrays. All 35 proteins showed considerable intratumoral heterogeneity within primary breast cancers with a mean coefficient of variation (CV) of 31% (range 22-43%). There were no significant differences between phosphorylated (CV 32%) and non-phosphorylated proteins (CV 31%) and in the extent of intratumoral heterogeneity within a defined tumor zone (CV 28%, range 18-38%) or between different tumor zones (CV 24%, range 17-38%). Lymph node metastases from the same patient showed a similar heterogeneity in protein expression (CV 27%, range 18-34%). In comparison, the variation amongst different patients was higher in primary tumors (CV 51%, range 29-98%) and lymph node metastases (CV 65%, range 40-146%). Several proteins showed significant differential expression between different tumor stages, grades, histological subtypes and hormone receptor status. Commonly used protein biomarkers of breast cancer, including proteins from HER2, uPA/PAI-1 and EGFR signaling pathways showed higher than previously reported intratumoral heterogeneity of expression levels both within primary breast cancers and between lymph node metastases from the same patient. Assessment of proteins as diagnostic or prognostic markers may require tumor sampling in several distinct locations to avoid sampling bias.     

Availability of dissolved organic carbon for planktonic bacteria in oligotrophic lakes of differing humic content

Bacterioplankton from 10 oligotrophic lakes, representing a gradient from clearwater to polyhumic, were grown in dilution cultures of sterile filtered lake water. The bacterial biomass achieved in the stationary phase of the dilution cultures was positively correlated with the amount of both humic matter and dissolved organic carbon (DOC) in the lakes. About the same fraction of the total DOC pool was consumed in the dilution cultures of all lakes (average 9.5%, coefficient of variation (CV) 24%), with approximately the same growth efficiency (average 26%, CV 28%). Thus, humic lakes could support a higher bacterial biomass than clearwater lakes due to their larger DOC pools. The relevance of the results to planktonic food webs of humic and clearwater lakes is discussed.     

Comparison of classical Lowry, modified Lowry and a dye-binding assay for the estimation of protein in allergen extracts and influence of different parameters on the modified Lowry assay

Protein values of dialysed allergen extracts determined by Lowry, modified Lowry (trichloroacetic acid precipitation of the proteins) and dye-binding assay were compared. The influence of different parameters on the modified Lowry was examined. The reproducibility of the modified Lowry was checked with three independent measurements. For the examination of recovery a constant amount of 6-grass pollen allergen proteins was added to the samples of the standardized human serum albumin prepared for the calibration curve. The samples were measured by modified Lowry. The mean of the ratio between the protein values of the dialysed allergen extracts obtained by modified Lowry and those obtained by classical Lowry was 3.59 (coefficient of variation Cv = 45%). The mean of the ratio between the protein values of the allergen extracts obtained by modified Lowry and dye-binding assay was 1:0.71 (Cv = 31%). Phenol interfered with the modified Lowry. Phenolic allergen extracts showed higher "protein values" than non-phenolic allergen extracts. This influence could be reduced by a second precipitation of the dissolved precipitate. The precipitation of non-phenolic dialysed aqueous allergen extracts was complete after the first trichloroacetic acid precipitation. By incubating samples with the Folin-Ciocalteu's reagent at 55 degrees C in a waterbath, the time necessary for developing the colour could be reduced from 45 min to 5 min. Protein measurements by modified Lowry of a 6-grass pollen allergen extract in three different laboratories showed good reproducibility. For these extract 785 micrograms protein/ml (Cv = 4%) could be measured.(ABSTRACT TRUNCATED AT 250 WORDS)     

Arbuscular mycorrhizae formed by Penicillium pinophilum improve the growth, nutrient uptake and photosynthesis of strawberry with two inoculum-types

Penicillium pinophilum was isolated from the soil in a commercial strawberry field. The strain readily formed arbuscular mycorrhizae (AM) with the roots of strawberry 'Zoji' (Fragaria x ananassa Duch. CV.) when plants were inoculated with either fresh cultured hyphae or root/soil mixtures. Fresh hyphae, however, resulted in higher amounts of colonization than root/soil inoculum. Compared with uninoculated strawberries, inoculation increased plant dry weight by 31%, as well as nitrogen content (47%), phosphorus content (57%), and photosynthetic rate (71%). AM inoculation also shortened the blossom and ripening date by 3 and 4 days, respectively. This is the first report of a P. pinophilum strain resulting in mycorrhiza with strawberry roots. The significant advantages of this strain are that it is easy to culture and inoculation of plants results in significant growth benefits that may be useful in strawberry production.     

Forage quality and quantity affect red mason bees and honeybees differently in flowers of strawberry varieties

Nectar is a vital source of energy for bees and other pollinators and pollen represents the only source of protein in the diet of bees. Nectar and pollen quality and quantity can therefore affect foraging choices. Strawberry, Fragaria × ananassa (Rosaceae), is a flowering crop that requires insect pollination for the berries to develop optimally. The solitary red mason bee, Osmia bicornis L. (Hymenoptera: Megachilidae), occurs naturally but like the eusocial western honeybee, Apis mellifera mellifera L. (Hymenoptera: Apidae), it is also a commercially reared pollinator used in strawberry production. We hypothesized that strawberry nectar and pollen quality would affect the foraging choice of these two types of bees. In this study nectar and pollen quality is represented by various levels of sugar and protein content, respectively, as well as the number of open strawberry flowers in the experimental field, would affect the foraging choice of these two types of bees. Consistent with previous studies, we found significant and major differences between strawberry varieties in proportions of sucrose in the nectar sugar and in pollen viability – a proxy for pollen protein content. All measured parameters had a significant effect on red mason bee visitation frequency. Contrary to expectations, honeybee foraging behavior was only affected by the number of open flowers and not by any of the quality parameters measured. Our findings indicate that red mason bees were capable of assessing nectar and pollen quality and prioritize accordingly. The pattern observed indicates that individual red mason bees changed foraging focus between strawberry varieties depending on whether nectar or pollen was collected. Our results suggest that targeted breeding of varieties toward high levels of nectar sugar and sucrose concentrations and high pollen protein content may increase pollination success from red mason bees and possibly other solitary bees.     

From the ground up: biotic and abiotic features that set the course from genes to ecosystems

Spatial variation in cone serotiny in Rocky Mountain lodgepole pine (Pinus contorta ssp. latifolia) across Yellowstone National Park influences initial pine recruitment after stand‐replacing fire with tremendous population, community, and ecosystem consequences. A previous study showed that much of the spatial variation in serotiny results from the balance of selection arising from high frequencies of fire favoring serotiny countered by opposing selection exerted by American red squirrels (Tamiasciurus hudsonicus) as seed predators. This earlier study, however, assumed stable local red squirrel densities over multiple generations of pines. Here, we examine environmental properties that might contribute to long‐term stability in the densities of red squirrels among sites. We found that the amount of clay in the soil, an indicator of plant and fungal growth—the latter an important food resource for red squirrels—and the coefficient of variation (CV) in diameter at breast height (DBH) of forest trees together account for a substantial amount of variation in red squirrel density. Soil development occurs over very long time scales, and thus, intersite variation in the amount of clay is unlikely to shift across pine generations. However, CV of DBH and squirrel density increase with stand age, which acts to amplify selection against serotiny with increasing interfire interval. Regardless, much of the variation in the CV of DBH is accounted for by soil bulk density, mean annual temperature, and surface curvature, which are unlikely to vary in their relative differences among sites over time. Consequently, these soil and abiotic attributes could contribute to consistent spatial patterns of red squirrel densities from one pine generation to the next, resulting in consistent local and spatial variation in selection exerted by red squirrels against serotiny.     

Nondiscrete heterogeneity of human erythrocytes: comparison of Coulter-principle flow cytometry and Soret-hemoglobinometry image analysis

In the blood of normal subjects, the volumes of single erythrocytes are distributed with a coefficient of variation (CV) of 10.8 +/- 1.8%; while in hemoglobinopathies, CV increases proportionately to the degree of anemia produced. Using single cell Soret-band hemoglobinometry and focused-aperture impedance counting, we compared the distribution of red cell volume, area, hemoglobin content, and hemoglobin concentration in normals and subjects with anemic disorders. The CV, nondiscrete heterogeneity, is first, a general characteristic of biologic measurement, second, a sensitive indicator of abnormality of erythropoiesis, and third, consistently less for hemoglobin concentration than for volume, area, or hemoglobin content of the same cells.     

Effects of cultivar, fruit number and reflected photosynthetically active radiation on Fragaria x ananassa productivity and fruit ellagic acid and ascorbic acid concentrations

BACKGROUND AND AIMS: A number of strawberry varieties were surveyed for their total ellagic acid concentration, and attempts were made to determine if ellagic acid and ascorbic acid concentrations of two strawberry cultivars could be increased by polythene reflective mulches. METHODS: After adjusting crop yields and cultivation using polythene mulches with two different PAR reflective capacities, field- and polytunnel-grown strawberries were analysed for ellagic acid and ascorbic acid concentrations by HPLC. Comparative measurements of yield and fruit quality were determined along with plant developmental changes. KEY RESULTS: Ellagic acid concentration varied widely with strawberry cultivar (60-341 microg g(-1) frozen weight), as did the ratio of conjugated ellagic acid : free ellagic acid. Also, there was significant year-to-year variation in total ellagic acid concentration with some cultivars. Mulches with different reflective capacities impacted on strawberry production; highly reflective mulches significantly increased growth and yield, the latter due to increases in fruit size and number. CONCLUSIONS: Highly reflective mulches significantly increased total concentrations of ellagic acid and ascorbic acid relative to control in fruit of different cultivars. The potential of agronomic practices to enhance the concentration and amounts of these important dietary bioactive compounds is discussed.     

Fermentation rates of grape juice: v. Biotin content of juice and its effect on alcoholic fermentation rate

Microbiological analysis showed that juices from white grapes had less biotin than juices from red grapes. The biotin content of the juices of some varieties was significantly different from that of other varieties. We did not note any regional effects on the biotin content of the juices. Biotin content of the Cabernet Sauvignon grapes increased significantly with maturity, whereas the biotin content of a white variety did not. The biotin content, with the total nitrogen, can be used to estimate indirectly the yeast growth potential and hence to predict the fermentation rate of the juice. About 84% of the rate variation can be accounted for by the calculated regression equations.     

LED光源对不同品种生菜生长和品质的影响

以自然光为对照(CK),探讨了LED红光(R)、蓝光(B)和红蓝组合光(RB)对不同品种生菜生长与品质的影响.结果显示:(1)不同光质处理的4个品种生菜的根长、株高及生物量积累等形态及生长指标具有相同的变化规律.(2)植株干鲜重、叶面积及根系活力在R和RB处理下都较大,而在B处理下较小;金祥、高华和永荣的B处理植株可溶性蛋白含量较高;联记、金祥和高华植株的淀粉含量在RB处理下较多,而永荣在R处理下较多;各品种植株可溶性糖含量在R和RB处理下较高,而永荣植株RB处理蔗糖含量较高,其余品种蔗糖含量在R处理下较高;金祥、高华和永荣VC含量在B和RB下较高,联记在RB下较高,各品种在R下均较低;植株总酚含量在各光质处理间无显著差异.(3)联记的硝态氮含量及亚硝酸还原酶对光质敏感,B处理能降低其硝态氮含量及亚硝酸还原酶活性,其他品种的硝态氮含量及亚硝酸还原酶活性在光质处理间无显著差异.研究表明,相同光质下品种间生长无显著差异,而各品种生菜植株在红光和红蓝组合光下生长较好,在红蓝光处理下品质较优,红蓝光是设施栽培生菜的良好光源.     

Lack of Detectable Allergenicity in Genetically Modified Maize Containing “Cry” Proteins as Compared to Native Maize Based on In Silico & In Vitro Analysis

BackgroundGenetically modified, (GM) crops with potential allergens must be evaluated for safety and endogenous IgE binding pattern compared to native variety, prior to market release.ObjectiveTo compare endogenous IgE binding proteins of three GM maize seeds containing Cry 1Ab,1Ac,1C transgenic proteins with non GM maize.MethodsAn integrated approach of in silico & in vitro methods was employed. Cry proteins were tested for presence of allergen sequence by FASTA in allergen databases. Biochemical assays for maize extracts were performed. Specific IgE (sIgE) and Immunoblot using food sensitized patients sera (n = 39) to non GM and GM maize antigens was performed.ResultsIn silico approaches, confirmed for non sequence similarity of stated transgenic proteins in allergen databases. An insignificant (p> 0.05) variation in protein content between GM and non GM maize was observed. Simulated Gastric Fluid (SGF) revealed reduced number of stable protein fractions in GM then non GM maize which might be due to shift of constituent protein expression. Specific IgE values from patients showed insignificant difference in non GM and GM maize extracts. Five maize sensitized cases, recognized same 7 protein fractions of 88-28 kD as IgE bindng in both GM and non-GM maize, signifying absence of variation. Four of the reported IgE binding proteins were also found to be stable by SGF.ConclusionCry proteins did not indicate any significant similarity of >35% in allergen databases. Immunoassays also did not identify appreciable differences in endogenous IgE binding in GM and non GM maize.     

Infectivity and effectiveness of Glomus intraradices on micropropagated plants

Colonization by Glomus intraradices takes place very early within the root system of micropropagated plantlets of strawberry (var. avanta, elsanta), raspberry (var. himboqueen, Zeva I), and hortensia (var. leuchtfeuer). The arbuscular mycorrhizal fungus (AMF) did not colonize roots of the different hosts to the same extent, and considerable differences were observed between the varieties. The results reported here confirm that endomycorrhizal root colonization is affected by the host-fungus combination. The effects ranged from mutualistic (hortensia), through neutral (strawberry var. avanta, raspberry var. Zeva I) to negative (raspberry var. himboqueen and strawberry var. elsanta). Non-mycorrhized (control) plants of strawberry produced more runners than mycorrhized plants under controlled growth conditions (phytotron). Transfer of the potted plants to the field resulted in drastic alterations in overall growth and development within 4 weeks. Mycorrhized plants became healthy, and mycorrhized strawberry plants produced many stout runners. The number of the runners and their biomass were almost the same (var. avanta) and treated plants produced even more runners than the controls (var. elsanta). The authors have demonstrated the need to determine the specific effects of each species of AMF on individual prospective host plants prior to their utilization in the micropagation of plantlets.     

Flower Volatiles,Crop Varieties and Bee Responses

Pollination contributes to an estimated one third of global food production, through both the improvement of the yield and the quality of crops. Volatile compounds emitted by crop flowers mediate plant-pollinator interactions, but differences between crop varieties are still little explored. We investigated whether the visitation of crop flowers is determined by variety-specific flower volatiles using strawberry varieties (Fragaria x ananassa Duchesne) and how this affects the pollination services of the wild bee Osmia bicornis L. Flower volatile compounds of three strawberry varieties were measured via headspace collection. Gas chromatography showed that the three strawberry varieties produced the same volatile compounds but with quantitative differences of the total amount of volatiles and between distinct compounds. Electroantennographic recordings showed that inexperienced females of Osmia bicornis had higher antennal responses to all volatile compounds than to controls of air and paraffin oil, however responses differed between compounds. The variety Sonata was found to emit a total higher level of volatiles and also higher levels of most of the compounds that evoked antennal responses compared with the other varieties Honeoye and Darselect. Sonata also received more flower visits from Osmia bicornis females under field conditions, compared with Honeoye. Our results suggest that differences in the emission of flower volatile compounds among strawberry varieties mediate their attractiveness to females of Osmia bicornis. Since quality and quantity of marketable fruits depend on optimal pollination, a better understanding of the role of flower volatiles in crop production is required and should be considered more closely in crop-variety breeding.     

A Sensitive Radial Diffusion Method for Detection of Esterolytic Activities of Proteases

The influence of under-nutrition (sub-maintenance feeding) and ad libitum feeding on the deposition of proteins in different subcellular sarcoplasmic fractions of red (tonic) and white (phasic) muscles of growing broilers was investigated. The relative concentration of overall sarcoplasmic proteins was lesser in red than in white muscles from ad libitum fed broilers. The content of mitochondrial proteins was slightly more and that of lysosomal and microsomal proteins and of true soluble proteins was lesser in red than in white muscles. Besides, the relative amount of some specific molecular weight proteins in each subcellular fractions differed by more than 50% between red and white muscle.

There was also conspicuous differences in the influence of under-nutrition on the proteins in red and white muscles. Some reduction in mitochondrial, lysosomal, microsomal, and soluble protein content occurred only in white muscle, whereas little change was found in subcellular fractions in red muscle from underfed broilers. The relative amount of some proteins in each subcellular fraction of both muscles remained unaffected, and that of others either decreased or increased more than 20 to 50% due to nutritional stress.     

Variability of the healthy human proteome

The aim of this review is to analyze results of studies on characteristics of protein variability and diversity of posttranslational modifications of proteins in healthy humans. Numerous studies have demonstrated that a proteomic profile is characterized by significant intra- and inter-individual variability, and quite often natural (“normal”) variability of some proteins can be comparable to changes observed in pathological processes. Results obtained by our research group have demonstrated high intra-individual variability of serum low-molecular subproteome of healthy volunteers, certified by a special medial committee (the coefficient of variation (CV) of 42.6%). The proteins characterized by high variability under normal conditions (e.g. haptoglobin — 0–40 mg/ml; lysozyme — 0.01–0.1 mg/ml; C-reactive protein — 0.01–0.3 mg/ml) cannot be considered as potential biomarkers of diseases. On the contrary, proteins and peptides characterized by insignificant dispersion in healthy population (such as albumin ( CV = 9%); transferrin-(CV = 14%); C3c complement (CV = 17%), α-1 acid glycoprotein (CV = 21%), α-2-macroglobulin (CV = 20%); transthyretin fragment (CV = 28.3%) and α2-HS-glycoprotein βchain (CV = 29.7%)) can provide valuable information about the state of health. Thus, studies of plasticity in the proteomic profiles of healthy humans will help to correct reference intervals used in clinical proteomics.     

Pollen allergens from different Dactylis glomerata varieties harvested between 1986 and 1988

Pollen was collected from different cultivated varieties of Dactylis glomerata in 1986, 1987 and 1988 and compared with an undifferentiated stock of Dactylis glomerata pollen harvested and stored dry at +4°C since 1981. The allergen content of the crude pollen extracts was established on the basis of the IgE antibodies from the sera of three different patients allergic to grass pollen, using the nitrocellulose immunoprint technique following isoelectric focusing (IEF) and separation in agarose gels. Coomassie blue and silver staining patterns were also compared. They showed some marked differences in the isoelectric points of the constituants of these extracts. Using this major allergen recognition by patient sera as a selection criteria we were able to delimit 8 Dactylis varieties with low or undetectable Dac g IV allergen amounts and 7 varieties which contained this allergen. Two other allergens used as markers enabled us to suggest a kind of taxonomy, based upon the allergen presence, of these 15 varieties. The effect of the storage temperature was studied for 8 different varieties kept at +4°C, -20°C and -40°C. The allergen recognition and silver staining patterns after IEF separation of the crude pollen extracts revealed a selective persistance of some constituants and the disappearance of others at +4°C. The effect of freeze-drying was also analysed in the same way. Finally the pollen constituants of one Dactylis glomerata variety, harvested in 1986 and 1987 and kept at different temperatures were compared using the same techniques.     

Changes in peroxidase activities and soluble proteins in strawberry varieties under salt-stress

Effects of salt stress on the activity of peroxidase (PRX) isozyme and leaf proteins were studied in the three strawberry varieties, Camarosa, Tioga and Chandler. Plants were watered with modified 1/3 Hoagland nutrient solution containing 0 (control), 8.5, 17.0 and 34.0 mM NaCl for 30 days fallowing 20 days acclimation. PRX activity was assayed in leaf extracts and PRX profiles of control and salt treated plants were performed by native polyacrylamide gel electrophoresis (PAGE). During the salt stress Tioga and Camarosa tolerated the cellular damage with less electrolyte leakage while Chandler exhibited a severe cellular damage with the highest (87.5 % in 34.0 mM NaCl treatment) electrolyte leakage. Total soluble protein content was decreased by salinity in Tioga (from 11.35 to 5.86 mg g⁻¹ DW) and Chandler (from 9.35 to 3.90 mg g⁻¹ DW) while it was almost unchanged in Camarosa. Salt stress increased total and specific PRX activity as compared with the control in all the varieties. In native PAGE, one basic isoperoxidase band (Rf=0.27) was observed commonly with different band intensity in all the treatments. Therefore, it might be associated with lignification and recovery of cell membrane damage in strawberry plants under salt stress.     

草莓果实发育成熟过程中糖苷酶和纤维素酶活性及细胞壁组成变化

以丰香和红丰草莓为试材,对果实发育成熟过程中细胞壁水解酶活性和细胞壁成份变化进行了研究．结果表明：半乳糖苷酶和α-甘露糖苷酶活性随草莓果实成熟而提高,葡萄糖苷酶活性不随草莓果实成熟而提高．随着果实发育成熟,纤维素酶活性、果胶酶活性不断提高．果实中未检测到内切多聚半乳糖醛酸酶活性,外切多聚半乳糖醛酸酶活性变化不随果实成熟软化而提高．随果实发育成熟,细胞壁中可溶性果胶和半纤维素增加,而离子结合果胶和共价结合果胶及纤维素减少．