套袋红富士苹果发育过程中果皮超微结构变化及相关酶类研究
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摘要
本试验以套袋红富士为试材，对其发育过程中果皮可溶性固形物、硬度、果皮粗纤
维、果皮果胶及与果皮细胞壁成分相关的酶活性进行了测定，并用透射电子显微镜对套
袋果和未套袋果发育过程中果皮细胞超微结构进行了系统观察。研究结果如下：
1.套袋后2个月内果实硬度无变化，从第3月开始下降，到采收期套袋果与对照果硬
度相差2.90 kg/cm2。套袋果可溶性固形物一直呈上升趋势，但始终低于对照果，且在套
袋后第1、3月上升平缓，在第2、4月上升较快，到采收期套袋较对照相差1.88个百分点。
2.套袋果和对照果粗纤维含量在套袋后前2个月呈下降趋势，在后2个月呈上升趋
势，且套袋果粗纤维含量一直低于对照果。二者原果胶变化基本相同，但套袋果原果胶
始终低于对照。套袋果可溶性果胶一直高于对照，套袋后1个月都呈上升趋势，套袋后2
个月都呈下降趋势。
3.套袋果的多聚半乳糖醛酸（PG）活性基本呈线性上升，且一直高于对照，在采收
期二者相差0.04个百分点。对照果和套袋果纤维素酶（Cx）活性在整个生长过程中都是
从低到高变化，变化幅度基本相同，且前者一直高于后者。套袋果多酚氧化酶（PPO）
活性一直低于对照果，在前期都呈上升趋势，在中期、后期都呈下降趋势。套袋果过氧
化物酶（POD）活性始终低于对照果，前期呈上升趋势，后期都呈下降趋势。
4.套袋与对照果皮细胞壁结构在果实生长前期致密，电子密度高，看不到细胞壁的
纤维素微纤丝，中胶层、次生壁和初生壁也不能区别开；在果实生长中期，套袋果细胞
壁开始变得松散，而对照果的仍较致密；在果实生长后期，套袋果细胞壁开始大量分解，
出现质壁分离现象，而对照果的才开始松散。
5.套袋果叶绿体在果实生长前期以前体存在，整个基质空间被大淀粉粒占据，而对
照叶绿体前体发达。在果实生长中期，套袋果叶绿体结构基本形成，只是其片层不发达，
而对照果的则完全形成，且片层很发达。到采收期套袋果叶绿体由于除袋后见光，在短
时间内已从潜在的质体中转化形成完整的片层结构，而对照果叶绿体减少，可能是参与
花青苷合成转化。
6.套袋果线粒体在整个果实生长过程中都较丰富、发达，采收期脂质小球丰富，但
中央大液泡漫布整个细胞，嗜锇颗粒大量存在。对照果的线粒体、脂质小球相对较少。
The main study contents of the thesis were the changing of the fruit hardness, the soluble
solid matter, the pericarp pectin and the coarse fibre and the activity of the enzymes involved the
pericarp cell wall by using Red‘Fuji’apple as material during apple development. At the same
time, the thesis observed the systemic ultrastructure changing of the pericarp of the bagged and
unbagged apple with electron microscope. The mainly study results including:
1. The fruit hardness was unchanged in two months after bagging, decreased after three
months, the hardness discrepancy of the bagged and unbagged apple was 2.90 kg/cm2 at the
harvesting time. The soluble solid matter of the bagged apple was still increasing, but always less
than than the unbagged apple, and increasing gently at the first and third month after bagging, and
increasing relatively quick at the second and fourth month. The soluble solid matter discrepancy
of the bagged and unbagged apple was 1.88 percent at the harvesting time.
2. The coarse fibre always decreased in former two months after bagging, increased at the
third and fourth months, and the coarse fibre of the bagged apple was still less than the unbagged
apple. The original pectin of the bagged and unbagged apple was approximately same, however
the content of the bagged apple was always less than that of the unbagged. The soluble pectin of
the bagged apple was still less than that of the unbagged, the content increased at first month,
decreased in two month after bagging.
3. The polygalacturonase activity of the bagged apple increased linearly, and always more
than that of the unbagged apple, and the discrepancy of the bagged and unbagged apple was
0.04 percent at the harvesting time. The cellulose enzyme activity of the bagged and
unbagged apple changed from low to high, and the changing extent was almost same, the
fomer was still higher than the after. The polyphenolo xidase activity of the bagged apple was
lower than that of the unbagged apple, and increased at the former period, decrease at the
middle and after period. The eperoxldase activity of the bagged apple was always lower than
that of the unbagged apple, increased at the former period, and decreased at the after period.
4. The pericarp cell wall structure of the bagged and unbagged apple was compact at the
former period of the fruit growth, the electron density was high, the cellulose micro filament
was invisible, the middle glue layer, the next and first cell wall were not distinguished; The
cell wall of the bagged apple began relaxing at the middle period of fruit growth, however the
unbagged apple’s was still relatively compact; The mostly cell wall of the bagged apple began
decomposing, and Incipient plasrmlysis appeared, whereas the unbagged apple's just began
relaxing.
5. The chloroplast of the bagged and unbagged apple existed with the orginal chloroplast
form at the former period of fruit growth, and its all space was held by the large starch grain,
however the unbagged apple's was development. The chloroplast structure of the bagged
apple was primarily formed at the middle period of fruit growth, just the segment
layer structure was not development, whereas that of the unbagged apple was already intact.
Because the bagged apples were expose to the sun after removing the paper bag, the potential
plastid changed into the intact segment layer structure at the short time, however that of the
unbagged apple was largely decreasing, maybe it participate in the anabolism and
transforming.
6. The mitochondrion of the bagged apple was abundance and development during all
course of fruit growth, the lipid bodies were richness, the central big vacuole besprinkled the
all cell, the osmiophilic globules abounded at the harvesting time, whereas the mitochondrion
and the lipid body of the unbagged apple were relatively less.
维、果皮果胶及与果皮细胞壁成分相关的酶活性进行了测定，并用透射电子显微镜对套
袋果和未套袋果发育过程中果皮细胞超微结构进行了系统观察。研究结果如下：
1.套袋后2个月内果实硬度无变化，从第3月开始下降，到采收期套袋果与对照果硬
度相差2.90 kg/cm2。套袋果可溶性固形物一直呈上升趋势，但始终低于对照果，且在套
袋后第1、3月上升平缓，在第2、4月上升较快，到采收期套袋较对照相差1.88个百分点。
2.套袋果和对照果粗纤维含量在套袋后前2个月呈下降趋势，在后2个月呈上升趋
势，且套袋果粗纤维含量一直低于对照果。二者原果胶变化基本相同，但套袋果原果胶
始终低于对照。套袋果可溶性果胶一直高于对照，套袋后1个月都呈上升趋势，套袋后2
个月都呈下降趋势。
3.套袋果的多聚半乳糖醛酸（PG）活性基本呈线性上升，且一直高于对照，在采收
期二者相差0.04个百分点。对照果和套袋果纤维素酶（Cx）活性在整个生长过程中都是
从低到高变化，变化幅度基本相同，且前者一直高于后者。套袋果多酚氧化酶（PPO）
活性一直低于对照果，在前期都呈上升趋势，在中期、后期都呈下降趋势。套袋果过氧
化物酶（POD）活性始终低于对照果，前期呈上升趋势，后期都呈下降趋势。
4.套袋与对照果皮细胞壁结构在果实生长前期致密，电子密度高，看不到细胞壁的
纤维素微纤丝，中胶层、次生壁和初生壁也不能区别开；在果实生长中期，套袋果细胞
壁开始变得松散，而对照果的仍较致密；在果实生长后期，套袋果细胞壁开始大量分解，
出现质壁分离现象，而对照果的才开始松散。
5.套袋果叶绿体在果实生长前期以前体存在，整个基质空间被大淀粉粒占据，而对
照叶绿体前体发达。在果实生长中期，套袋果叶绿体结构基本形成，只是其片层不发达，
而对照果的则完全形成，且片层很发达。到采收期套袋果叶绿体由于除袋后见光，在短
时间内已从潜在的质体中转化形成完整的片层结构，而对照果叶绿体减少，可能是参与
花青苷合成转化。
6.套袋果线粒体在整个果实生长过程中都较丰富、发达，采收期脂质小球丰富，但
中央大液泡漫布整个细胞，嗜锇颗粒大量存在。对照果的线粒体、脂质小球相对较少。
The main study contents of the thesis were the changing of the fruit hardness, the soluble
solid matter, the pericarp pectin and the coarse fibre and the activity of the enzymes involved the
pericarp cell wall by using Red‘Fuji’apple as material during apple development. At the same
time, the thesis observed the systemic ultrastructure changing of the pericarp of the bagged and
unbagged apple with electron microscope. The mainly study results including:
1. The fruit hardness was unchanged in two months after bagging, decreased after three
months, the hardness discrepancy of the bagged and unbagged apple was 2.90 kg/cm2 at the
harvesting time. The soluble solid matter of the bagged apple was still increasing, but always less
than than the unbagged apple, and increasing gently at the first and third month after bagging, and
increasing relatively quick at the second and fourth month. The soluble solid matter discrepancy
of the bagged and unbagged apple was 1.88 percent at the harvesting time.
2. The coarse fibre always decreased in former two months after bagging, increased at the
third and fourth months, and the coarse fibre of the bagged apple was still less than the unbagged
apple. The original pectin of the bagged and unbagged apple was approximately same, however
the content of the bagged apple was always less than that of the unbagged. The soluble pectin of
the bagged apple was still less than that of the unbagged, the content increased at first month,
decreased in two month after bagging.
3. The polygalacturonase activity of the bagged apple increased linearly, and always more
than that of the unbagged apple, and the discrepancy of the bagged and unbagged apple was
0.04 percent at the harvesting time. The cellulose enzyme activity of the bagged and
unbagged apple changed from low to high, and the changing extent was almost same, the
fomer was still higher than the after. The polyphenolo xidase activity of the bagged apple was
lower than that of the unbagged apple, and increased at the former period, decrease at the
middle and after period. The eperoxldase activity of the bagged apple was always lower than
that of the unbagged apple, increased at the former period, and decreased at the after period.
4. The pericarp cell wall structure of the bagged and unbagged apple was compact at the
former period of the fruit growth, the electron density was high, the cellulose micro filament
was invisible, the middle glue layer, the next and first cell wall were not distinguished; The
cell wall of the bagged apple began relaxing at the middle period of fruit growth, however the
unbagged apple’s was still relatively compact; The mostly cell wall of the bagged apple began
decomposing, and Incipient plasrmlysis appeared, whereas the unbagged apple's just began
relaxing.
5. The chloroplast of the bagged and unbagged apple existed with the orginal chloroplast
form at the former period of fruit growth, and its all space was held by the large starch grain,
however the unbagged apple's was development. The chloroplast structure of the bagged
apple was primarily formed at the middle period of fruit growth, just the segment
layer structure was not development, whereas that of the unbagged apple was already intact.
Because the bagged apples were expose to the sun after removing the paper bag, the potential
plastid changed into the intact segment layer structure at the short time, however that of the
unbagged apple was largely decreasing, maybe it participate in the anabolism and
transforming.
6. The mitochondrion of the bagged apple was abundance and development during all
course of fruit growth, the lipid bodies were richness, the central big vacuole besprinkled the
all cell, the osmiophilic globules abounded at the harvesting time, whereas the mitochondrion
and the lipid body of the unbagged apple were relatively less.
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