Differences in ovary size among olive (Olea europaea L.) cultivars are mainly related to cell number,not to cell size

Differences in mature fruit size among olive cultivars are related to differences in ovary size at bloom, but it is not known whether cell number or size determines the variation in ovary size. In this study we measured cell size and number in equatorial cross-sectional areas of the principal ovary tissues (mesocarp and endocarp) in eight cultivars with very different fruit and ovary size. The results showed that cell number explained most of the differences in ovary size, while cell size was not related to ovary size, except for a weak (R² = 0.33) correlation in the endocarp. Since sink strength is thought to be related to cell number, this finding supports the hypothesis that larger ovaries represent stronger sinks. The implications of greater sink strength with bigger ovaries are discussed. Within cultivars, while the cross-sectional areas of mesocarp and endocarp were similar, cell number was higher in the mesocarp but cell size was smaller compared to the endocarp.     

Early growth and development of the olive fruit mesocarp

SUMMARY

We examined growth and development of the ‘Manzanilla’ olive fruit mesocarp in transverse equatorial sections during the first 12 weeks after full bloom (AFB). Sequential sampling and quantitative data provided an integrated view of the formation of this tissue. The mesocarp, or fruit flesh, was formed by relatively isodiametric parenchyma cells with a small number of isolated sclereids. By four weeks AFB a gradient in cell size characteristic of mature olive fruits began to appear. Biweekly measurements of cell size and number indicated that, as in other drupes, both cell division and expansion contribute to initial mesocarp growth. From six weeks AFB, further mesocarp growth was determined solely by cell expansion. Transverse areas of mesocarp and endocarp, also measured biweekly, revealed that both tissues expand in a similar manner until eight weeks AFB, after which mesocarp growth predominated.     

Olive fruit pulp and pit growth under differing nutrient supply

The objetive of this work was to study if the addition of nutrients to the irrigation water modified ‘Manzanilla de Sevilla’ olive pulp and pit growth. The experiment was carried out during the 2003 fruit-growth period in an irrigated orchard near Seville, southern Spain. Fruit samples were taken in July and September, at 12 and 21 weeks after full bloom (AFB) respectively, in trees irrigated with (T1) or without (T0) the addition of nutrients (N–P–K). The nutrient availability of T1 fruits increased the fruit fresh and dry weight, longitudinal and equatorial diameters, and the pulp-to-pit ratio, characteristics particularly appreciated for table olives. The balance of growth between the fruit mesocarp (pulp) and endocarp (pit) was modified because those two tissues were affected differently. Mesocarp fresh weight was significantly higher at both 12 and 21 weeks AFB in the fertilized treatment, as was mesocarp dry weight at 12 weeks AFB. Neither the endocarp fresh and dry weight nor shape (the ratio of the equatorial and longitudinal diameters) was altered at either of the two studied dates. These results emphasize the importance of an appropriate fertilization management in irrigated olive orchards, particularly for table olives, and also confirm the olive endocarp as a strong sink tissue that competes with the mesocarp during early development.     

Fruit set is inversely related to flower and fruit weight in olive (Olea europaea L.)

Fruit weight is strongly correlated to ovary weight in olive as big-fruited cultivars have larger ovaries and flowers at bloom. We tested the hypothesis that larger ovaries imply stronger sinks and therefore reduce fruit set, expressed as number of fruits. Flower/fruit load per centimeter of shoot was assessed every month from flowering to harvest, on sample shoots of many different olive cultivars differing in fruit size. Data were taken for two years: 2007 and 2008. Additionally, in 2008, a fruit thinning experiment was carried out by leaving a single fruit per shoot, to assess the maximum potential fruit weight achievable by each cultivar. Results indicated that fruit size was mostly genetically determined as thinned fruits of small-fruited cultivars, although bigger than control fruits, never got as big as fruits from large-fruited cultivars, confirming that large ovaries are a prerequisite for large fruits. Among all cultivars, the number of flowers per inflorescence, the number of inflorescences, and the number of flowers per centimeter of shoot were not correlated to average flower weight while total flower mass per centimeter of shoot was positively correlated to average flower weight (i.e. large flowered/fruited cultivars had greater total flower mass). However, one month after bloom and thereafter, all parameters were negatively and exponentially correlated with fruit average weight across cultivars, except fruit mass per centimeter of shoot which was generally not correlated to fruit average weight, except in one year, at harvest, when fruit mass per centimeter of shoot was positively correlated to fruit average weight. These results suggest that fruit set is a consequence of, and inversely proportional to flower/fruit size in olive.     

Fruit weight is related to ovary weight in olive (Olea europaea L.)

Fruit size is an important parameter both for scientific understanding and for commercial purposes. In many species, mature fruit size is often related to floral ovary size, but no literature exists in olive that demonstrates such a relationship. Previous work suggests that olive cultivars with different fruit sizes have similar cell number and size in the ovary transectional area, but ovary and fruit dry weight was not measured. In the present study, ovary dry weight and fruit dry weight during the whole fruit development season until harvest were measured in olive cultivars with different fruit size, over three years. Flower dry weight was also measured. Fruit weight at harvest was strongly correlated to ovary weight at bloom, both in single-year data and when data from three years were pooled. Flower dry weight, excluding the ovary, was also correlated to ovary dry weight. Ovary dry weight was strongly correlated not only to the fruit dry weight at maturity, but also at any date during fruit development. The mature fruit/ovary dry weight ratio ranged between 1000 and 4000 among cultivars, but was not correlated to the fruit dry weight at maturity. These results suggest that, in olive, fruit weight is genetically controlled through the ovary weight at bloom. This knowledge may have implications in the understanding of fruit set and source-sink relationships in olive.     

The relationship between fruit size and cell division and enlargement in cultivated and wild persimmons

Summary

To determine the anatomical basis for differences in fruit size in Japanese persimmon (Diospyros kaki Thunb.), the number and sizes of parenchymal cells were measured in four astringent cultivars and in wild D. lotus over a single season in Japan. Fruit weight at harvest correlated with the final number of parenchymal cells (r = 0.95*) and their size (r = 0.92*). There was also a correlation between fruit diameter and the number of cell layers at full bloom (r = 0.93*), which increased 1.7 – 1.9-fold during fruit development.The duration of cell division and the sizes of the cells at full bloom were similar among the cultivars, whereas mean parenchymal cell lengths increased by 7.7-, 6.7-, 5.4-, 5.0- and 3.8-fold from full bloom to harvest in ‘Otanenashi’ (average fruit diameter = 96 mm), ‘Hiratanenashi’ (77 mm), ‘Saijo’ (54 mm), ‘Tsurunoko’ (44 mm), and D. lotus (20 mm), respectively. The final size of tannin cells was larger in ‘Otanenashi’ and ‘Hiratanenashi’ than in the other cultivars, including the wild species.These results indicate that the size of fruit at harvest was determined by the number of cells at full bloom, and their expansion during fruit development.     

龙眼果皮发育解剖学观察

以储良龙眼品种为试材,观察了果皮的解剖结构发育特点。果皮发育早期(花后10d前),细胞体积小,排列紧密,细胞分裂活跃,外果皮有凸起的皱褶结构,有大量的表皮毛分化。果皮发育中期(花后10～52d),细胞体积变大,表皮毛开始脱落,外果皮逐渐平滑,中果皮处有石细胞群和海绵状组织的分化。果实发育后期(花后52d后,假种皮快速膨大阶段),有木栓形成层出现,形成周皮代替表皮起保护作用,随着果实生长,木栓层出现局部破裂。龙眼果皮可划分为外、中、内3层。中外果皮在发生次生木栓化前,由外表皮细胞及其附属物和角质层组成;在形成木栓形成层产生次生结构后,外果皮则由周皮组成。内果皮来源于子房内壁的几层细胞,由一层内表皮细胞和与之紧密相连的几层薄壁细胞组成。中果皮根据其组织特点可进一步分为上中果皮和下中果皮,上中果皮包括石细胞群、外层维管束和外果皮之内的薄壁细胞;下中果皮则主要包括海绵组织和其中的维管束。     

疏花对大久保桃中果皮细胞分裂与膨大的影响

 研究了大久保桃树〔Prunus persica (L. ) Batsch. ‘Okubo’〕疏花后果实生长发育期间果皮细胞分裂与膨大的变化。结果表明: 果皮细胞分裂一直持续到盛花后6周。疏花明显促进了花后3周内幼果中果皮的细胞分裂, 增加了果实中果皮细胞层数和果皮厚度, 这是导致成熟果实体积增大的主要原因。     

Improving ‘Bing’ sweet cherry fruit quality with plant growth regulators

Final fruit diameter is the prime determinant of sweet cherry fruit value. Previous research has shown that mesocarp cell size accounts predominantly for variability in final fruit size, within a genotype. Our research program evaluated the potential to improve sweet cherry fruit size/weight with growth regulators to affect cell division and/or cell expansion stages. In the current study we screened 8 plant growth regulators (PGRs), including cytokinins, gibberellins, and auxins, and their combinations for their ability to increase ‘Bing’ fruit weight. Each PGR was mixed in lanolin paste and applied to fruit pedicels at 9 or 30 days after full bloom (DAFB), to coincide with estimated peak in cell division and cell expansion activity, respectively. Several cytokinins applied 30 DAFB improved fruit weight significantly (ca. +15%) with N-(2-Chloro-4-pyridyl)-N′-phenylurea (CPPU) and 6-(3-hydroxybenzylamino) purine (mt-Topolin) at 100 mg l⁻¹ being the most effective. Gibberellins, applied alone, improved fruit size and delayed fruit maturation and exocarp coloration. GA₃ at 200 mg l⁻¹ applied at 9 DAFB was the most effective and improved final fruit weight by 15%. Fifty-six percent of the fruit from this treatment were ≥9 g compared to 15% of similar weight fruit from untreated limbs. Both GA₃ and GA_4/7 treatments applied 9 DAFB increased fruit radial expansion. 4-Chlorophenoxyacetic acid, a synthetic auxin, also stimulated higher fruit growth rates at stage I and stage II, and fruit color development, but did not improve final fruit size.     

Cellulose synthesis during endocarp hardening of peach fruit

Summary

Changes in cellulose content and the accumulation of PpCesA1 mRNA, putatively encoding the catalytic subunit of cellulose synthase (CesA), were analysed in endocarp and mesocarp during endocarp hardening in peach fruit. The cellulose content of the endocarp was undetectable or very low [< 0.1 mg g^–1 fresh weight (FW) of endocarp] until 48 d after full bloom (DAFB), but began to increase rapidly at 55 DAFB, when the endocarp had attained its maximum size, and reached 4.4 mg g^–1 FW of endocarp at 85 DAFB. In contrast, the cellulose content of the mesocarp reached a detectable level (1.5 mg g^–1 FW of mesocarp) at 34 DAFB, stayed at this level until 85 DAFB, but then was lower at 107 DAFB. PpCesA1 mRNA was detected in the endocarp throughout the hardening, but was undetectable in the mesocarp. It is suggested that PpCesA1 has a role in the rapid synthesis of cellulose in the secondary cell walls of endocarp tissue during hardening.     

Developmental problems in over-winter off-season longan fruit. II: Development of pericarp structure

Fruit development was investigated in field and changes in pericarp structure were comparatively studied in over-winter off-season and on-season longan (Dimocarpus longan Lour. cv. Chuliang). The results showed developmental problems including small fruit size, severe fruit cracking and heavy fruit drop in the off-season fruit, although their seed size was not significantly affected compared to the on-season fruit. Anatomical studies showed that off-season longan fruit had a poorly developed pericarp, which was reflected by smaller fresh weight as well as thinner pericarp thickness, thinner spongy tissue, fewer cell layers in the periderm, smaller cell size and fewer cell numbers in the parenchyma tissue at the upper mesocarp, and fewer cell layers in the endocarp. In contrast, the recovery of the sclereids in the mesocarp was higher in the pericarp of the off-season fruit. The poor development of the pericarp in off-season fruit probably caused the small fruit size and severe fruit cracking. It was suggested that the adverse climatic conditions encountered by fruit development in the over-winter off-season longan suppressed cell division and expansion in the pericarp but did not affect sclereid formation.     

Pistil abortion is related to ovary mass in olive (Olea europaea L.)

Pistil abortion is considered to be an evolutionary adaptation to save resources in andromonoecious species, balancing the number of pistils with the resources available. It follows that large-fruited species/cultivars should have greater levels of pistil abortion. Working in olive, we found that pistil abortion, expressed as percentage of staminate flowers, was positively correlated to the average ovary mass at bloom, across cultivars with different ovary/fruit mass. Since ovary mass correlated with fruit mass, pistil abortion also correlated with fruit mass. The absolute number of perfect flowers per inflorescence was negatively correlated with both ovary mass and pistil abortion, while the number of staminate flowers per inflorescence increased with both parameters. The total number of flowers (i.e. staminate + perfect) was not correlated to ovary mass or pistil abortion, suggesting that male function was not affected. The results support the hypothesis that pistil abortion is related to competition for resources among ovaries, and suggest that genetic differences in pistil abortion among olive cultivars may be explained by their different pistil mass and sink strength.     

Commercial Iranian olive cultivars: morphological traits,molecular diversity,and genetic structure

Iran is considered to have a unique gene pool of different fruit and nut species including olive (Olea europaea L.). In this study, we used 22 previously developed microsatellite (simple sequence repeat; SSR) markers for olive to evaluate the level of genetic variation and to produce identification keys for 63 Iranian accessions of olive belonging to 17 groups of cultivars. Based on morphological features, the number of flowers per inflorescence, fruit weights, endocarp weights, oil percentages, and flesh weights per endocarp had the highest coefficient of variation values, indicating the large extent of morphological variability among the 63 Iranian olive cultivars studied. All 22 microsatellite (SSR) markers revealed a high level of polymorphism, with a mean polymorphic information content (PIC) value of 0.511. Analyses of genetic structure among the 63 olive accessions were carried out using model-based methods, which showed a tendency for geographical clustering. Ten SSRs out of the 22 were successful for the identification of unique ID keys for 52 of the 63 accessions. In most cases, there was disagreement between the molecular data and the morphological data. These results could be used to reconstruct and maintain a collection of olive for future breeding programmes.     

桃果实缝合线软化过程中内源激素的变化

为了明确桃果实缝合线过早软化的原因,以大久保桃为试材,利用气质联用和高效液相色谱的方法,对缝合线软化果(障碍果)和正常果的种仁、缝合线处果肉和其它部位果肉中的生长素(IAA)、玉米素(Z)、脱落酸(ABA)、茉莉酸(JA)的浓度进行了测定分析。结果表明,果肉ABA浓度是种仁的8～9倍,并随着果实的发育逐渐增加,接近成熟时达到最高,主要表现出对果肉成熟的调控作用。障碍果中Z的浓度在内果皮硬化初期明显高于正常果,与其缝合线异常生长(凸起)吻合;障碍果种仁的IAA浓度在内果皮硬化初期明显低于正常果,果肉中IAA浓度在内果皮硬化后上升,且障碍果高于正常果,说明IAA对中果皮细胞的扩大与成熟软化起调控作用。JA在种仁中的浓度是果肉的3～6倍,在硬核初期障碍果明显低于正常果,到硬核结束时则高于正常果,推测可能JA与内果皮发育有关。     

荔枝果皮发育细胞学研究

 以‘淮枝’荔枝为试材, 对花前子房和花后果皮发育的解剖学特点进行了观察。荔枝果皮可划分为内、中、外3 层, 外果皮由单层表皮细胞、龟裂片峰处表皮细胞上的角质、裂纹处表皮细胞上的薄壁细胞构成; 中果皮是构建果皮的主体, 由龟裂片峰下的厚壁组织、上中果皮和下中果皮构成; 内果皮由薄壁未木栓化的皮层细胞构成。果皮的细胞分裂主要发生在开花前, 花后不同部位的果皮细胞还有一定的分裂行为, 但分裂的旺盛程度和停止时间有所不同, 下中果皮停止早, 约在花后19 d , 上中果皮其次, 约在花后32 d , 内果皮和裂纹处外果皮最晚, 约在花后47 d。细胞膨大规律总体呈“慢- 快- 慢”的S 型。     

桃果实维管束的分布及解剖研究

 为了解桃果实输导组织分布及结构特征, 以‘大久保’桃果实为试材, 观察了维管束系统的分支体系和分布状况, 并用石蜡切片技术对维管束组织的解剖结构进行了系统观察。结果表明, 桃果实中的维管束分为骨干、分支和毛细3级, 其中骨干维管束又分为种胚维管束、内果皮维管束和中果皮维管束(腹维管束、背维管束及分布在内果皮两侧着生于果柄的维管束) , 分别为种仁、内果皮、中果皮的物质运输通道。种胚维管束韧皮部极其发达, 并伴有三生结构; 内果皮中分布的维管束韧皮部发达; 腹维管束、背维管束粗大且分支较多, 兼具发达的韧皮部及木质部。分支维管束和毛细微管束主要分布在中果皮中。毛细维管束的特点是韧皮部处有腔, 腔体随着果实的发育不断扩大。形态和结构上的不同是与其功能相适应的, 由此推测种仁和内果皮发育需要的营养物质以有机营养为主, 中果皮发育需要的营养物质则涵括有机物、无机物、水分。     

红肉桃果实发育过程中色素和糖酸含量的变化

以红肉、白肉和黄肉3种不同果肉颜色的12份桃品种为材料分析成熟时果肉中花色苷的含量变化,结果表明,红肉桃与白肉、黄肉品种间花色苷含量差异显著,且红肉桃品种间的花色苷含量差异也很大。利用3个不同果肉颜色的桃品种进一步研究果实发育过程中色素及其相关物质的变化,结果表明,红肉桃品种天津水蜜在盛花后25d果肉中花色苷含量较高,而后逐渐减少;直至进入盛花后45d才缓慢回升,并在果实发育后期(盛花后86d)含量迅速增加;与此对应,该品种果皮在幼果期(盛花后35～45d)和果实发育后期(盛花后86d)2次出现红色,但果肉直到盛花后86d才明显变红;在整个果实发育过程中,天津水蜜果肉其它内含物的变化与白肉桃品种豫白和黄肉桃品种NJC83相似:叶绿素含量逐渐减少;类胡萝卜素和可溶性糖不断增加;可滴定酸前期上下波动变化,但近成熟时天津水蜜的可滴定酸显著高于其它2个品种。     

Identifying the location of olive fruit abscission

For high quality oil, table olive characteristics, and reduction of alternate bearing tendencies, it is often desirable to harvest olive fruits prior to full physiological maturity, that is, before the natural separation zone has differentiated. Olive fruit abscission can occur at different positions, mainly fruit-pedicel, pedicel rachis and peduncle-branch, apparently varying according to cultivar, fruit weight, and fruit maturation. Precisely identifying the location of separation is critical for studying the differentiation of the abscission zone or zones and testing the effectiveness of harvest procedures, including the mode of action of different fruit-loosening chemicals. We determined the olive fruit abscission zone position under natural conditions, throughout the complete fruit maturation period, for two cultivars differing in fruit size. The major separation zone for the cultivars studied was between the fruit and pedicel. For cv. Picual, abscission in zones other that fruit-pedicel occurred in less than 15% of the fruit at all dates throughout the maturation period, whereas in cv. Hojiblanca, the percentages of fruits detached at peduncle-shoot (27%) and pedicel-rachis (19%) were initially substantial, and then decreased progressively during fruit maturation. Our data clearly indicate both varietal and temporal differences which should be taken into account not only when testing fruit-loosening compounds but in extrapolating test results to different varieties or dates. Fruit weight did not appear affect the abscission location. The method we used, of counting the different detached units, is simple and effective for determining the position of fruit abscission.