Cultivar-based fruit size in olive depends on different tissue and cellular processes throughout growth

In drupe fruits, in addition to fruit size, the proportions of mesocarp and endocarp tissues are critical objectives for fruit quality, crop production and management. The olive fruit is a typical drupe, with cultivars which show a wide range in both fruit size and the proportions of mesocarp and endocarp. Characterizing the roles of tissue and cellular processes in producing genetically based fruit size variability is necessary for crop improvement, as well as deepening our understanding of fruit developmental physiology. This study used microscope image analysis to evaluate cell number and size, the growth of mesocarp and endocarp tissues, and their developmental timing in producing fruit size among six olive cultivars with a large range of fruit size. We found that cultivar mesocarp and endocarp size increased linearly with fruit size, with larger sizes favoring an increasingly greater mesocarp/endocarp ratio. Within the mesocarp, cultivar-based fruit size related directly to cell number and was established soon after bloom by cell division rate. In spite of different cell division rates, all cultivars showed similar timing of cell division activity, with the majority of cells produced in the two months after bloom but, surprisingly, a substantial number of cells formed during the following 6 months. Cell expansion was high throughout fruit growth and an important factor in achieving final fruit size, but cell size did not differ among cultivars at any time. We can conclude that fruit size differences among olive cultivars are due at the tissue level to both mesocarp and endocarp sizes and at the cellular level to cell division throughout fruit growth. Furthermore, since cell size is consistent among cultivars in spite of variable cell division, it is likely that cultivar differences in cell expansion accompany those in cell division.     

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.     

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.     

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。     

Productivity of olive trees with different water status and crop load

SUMMARY

A field experiment was conducted over two growing seasons to determine the combined effect of crop load and irrigation on yield components of olive trees (Olea europaea L. ‘Leccino’) planted at 6 m 3.8 m in a sandy-clay soil. Different crop loads were established by manual thinning of fruits. Drip irrigation was managed to maintain pre-dawn leaf water-potentials (PLWP) within the following ranges: (i) higher than –1.1 MPa (FI; fully irrigated); (ii) between –1.0 and –3.3 MPa (DI; deficit irrigated); or (iii) below –1.2 MPa, but not lower than –4.2 MPa (SI; severe deficit irrigated). The irrigation period lasted from 6 – 16 weeks after full bloom (AFB) in 2003, and from 5 – 19 weeks AFB in 2004. In 2003, full bloom was on 26 May; in 2004, it was on 3 June. Neither irrigation regime nor crop load affected flowering or flower quality the following Spring. The combined fruit yields [on a fresh weight (FW) basis] over both years in SI and DI trees were 49.0% and 81.6% of FI trees, respectively. The oil yields of SI and DI trees were 52.5% and 81.2% of FI trees, respectively. Fruit FWs in FI trees were greater than those of DI or SI trees at 8 weeks AFB. At harvest, FI trees bore the largest fruits, and SI trees the smallest fruits. The FWs of individual fruits at harvest in the FI and DI treatments decreased as crop load increased, but no such relationship was apparent for SI trees. The oil content of the mesocarp increased as PLWP increased from approx. –3.5 MPa to –1.5 MPa. The oil content of FI trees at harvest decreased from 53.1% to 45.7% dry weight as fresh fruit yield increased from 5 – 25 kg dm^–2 trunk cross-sectional area. However, crop load did not have any effect on the oil content of the mesocarp in DI trees. Fruit maturation was delayed by irrigation. Maturation index also decreased (indicating delayed maturation) as the crop load on FI or DI trees increased, but did not vary with crop level in SI trees.     

Effects of drought on fruit set,yield and quality of custard apple (Annona spp. hybrid) ‘African Pride’ plants

Summary

Studies were conducted to evaluate the effects of drought during flowering for both container- and field-grown trees, and during fruit development for container-grown trees, on subsequent growth, flowering, yield and quality of the custard apple ‘African Pride’ in subtropical Australia. Moderate drought (ψ_L=–1.5 MPa) reduced shoot growth by 20–30% and increased the number of flowers per lateral by about 40% compared with well-watered controls due to reduced apical dominance and increased lateral branching. Overall fruit set was not adversely affected by drought. Drought also increased the number of fruit harvested per tree by 47% compared with well-watered controls. This response was mainly due to the increase in flower number per tree. Drought reduced average fruit size by 11% possibly due to effects on cell division in the first 4–6 weeks after fruit set or on net carbon assimilation. In summary, mild to moderate drought during the flowering period and fruit set increased flowering without adversely affecting fruit set, but decreased fruit size.     

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.     

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.     

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。     

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.     

The effects of storage temperature and fruit size on firmness,extractable juice,woolliness and browning in two nectarine cultivars

Fruit firmness, extractable juice, woolliness and browning of the mesocarp tissue in ‘Independence’ and ‘Flavortop’ nectarines stored at —0.5°, 3°, 5° and 7°C for four weeks were determined during ripening at 15°C. The firmness of both ‘Independence’ and ‘Flavortop’ during ripening decreased as storage temperatures increased. The percentage extractable juice after cold storage and during ripening varied considerably between cultivars and between the storage temperatures. The extractable juice of fruit stored at higher temperatures tended to increase during ripening, whereas fruit stored at lower temperatures tended to decrease first before increasing. At storage temperatures of —0.5° and 3°C both cultivars passed through a stage of woolliness during ripening, while less woolliness occurred after storage at 5° and 7°C. In both cultivars the percentage extractable juice during ripening was higher on average at storage temperatures of 5° and 7°C. Severe browning of mesocarp tissue in both cultivars occurred during ripening after storage at 3°C. The effect of fruit size on changes in firmness, development of woolliness and mesocarp browning in ‘Flavortop’ nectarines stored at — 0.5°C for four weeks and ripened at 15°C was also determined. Larger nectarines lost firmness more rapidly, woolliness occurred sooner and the mesocarp tissue was more prone to browning than smaller fruit during ripening.     

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.     

甜瓜果实发育的组织结构观察

采用石蜡切片技术对甜瓜果实发育的组织结构进行了观察。结果表明：甜瓜外果皮包括表皮、气孔器和表皮毛,在幼果生长过程中,外表皮细胞主要进行垂周分裂,以适应果实体积的增大,随着果实的逐步发育,表皮细胞间隙逐渐填充一些含有木质素成分的物质,并在表皮细胞表面形成一层较厚的角质层。中果皮分为上中果皮和下中果皮,上中果皮包括薄壁细胞和横向伸长细胞,下中果皮则主要包括海绵组织和其中的维管束。幼果期,中果皮细胞排列紧密,各部分细胞大小及形态无明显差异。随着果实的发育,细胞分裂速度减慢,细胞体积不断增大,中果皮细胞径向直径呈现出梯度变化,由外向内逐渐增大。在果实成熟过程中,下中果皮大部分细胞高度分散,细胞壁破裂,细胞互相融合。构成细胞壁的纤维素成分快速溶解,果实软化成熟。     

Effect of calcium supply on the physiology of fruit tissue in ‘Andesu’ netted melon

Summary

Fruit development and physiological traits were evaluated in ‘Andesu’ netted melon plants grown with and without calcium. Calcium exclusion accelerated softening, alcoholic fermentation and ethylene evolution of fruit compared with those supplied with calcium, but the opposite trend occurred with sucrose accumulation. A significant difference of calcium concentrations in the fruit between treatments was observed at the developing stage of the fruit (P<0.05), but the difference became smaller as fruit matured. In the mesocarp tissue from the basal hemisphere, calcium concentrations in the NaCl soluble fraction at ripe fruit stage differed significantly (P<0.05) between calcium treatments but no significant difference was detected in the inner mesocarp of the distal hemisphere. Calcium exclusion did not lead to the development of water-soaked symptoms in the mesocarp tissue of ripe fruit. Uronic acid concentrations in ionically bound pectin fraction did not differ between calcium treatments throughout fruit development. In covalently-bound pectin fraction, however, uronic acid concentration of ripe fruit was significantly lower in calcium-excluded plants than in calcium-supplied plants (P<0.05). Our results suggest that: first, accelerated fruit softening under calcium deficient conditions might result from promoted ethylene evolution rather than from the shortage of ionically-bound calcium in cell walls, second, calcium deficient condition does not necessarily lead to water-soaked symptoms in tissue of ‘Andesu’ melon fruit.     

Effects of Thinning Blossom and Fruitlets on Growth and Cropping of Sunset Apple

The results of a fruit-thinning trial, by hand, on the apple variety Sunset on M.IX are reported. Thinning flowers at the pink bud stage, and fruitlets at one, two and three weeks after full bloom, increased fruit size and trunk-girth increment. The two earliest treatments increased the number of shoots but mean shoot length was unaffected by any of the treatments.

The increase in fruit size was due to an increase in cell number per fruit rather than to an increase in cell volume. The concept of competition between meristematic regions of the aerial parts of the tree is discussed. No residual effect of thinning on yield, fruit size or shoot growth occurred in the season following the three-year experimental period, when the trees were left untreated.     

Gold speckles and crystals in tomato fruits (Lycopersicon esculentum Mill.)

Summary

Tomato fruit walls often possess tiny yellowish spots around the calyx and shoulders of the fruit, so-called gold speckles. In some cases these become very numerous and lead to down-grading of the fruit. In such fruit crystal sand is present in separate cells and groups of cells in the outermost parenchymatous tissue layers of the mesocarp. This crystal sand causes the gold speckles. Also raphid crystals occur in great quantities in the pulp of those tomatoes which display numerous gold speckles in the wall. Druse crystals are absent. Both crystal sand and raphid crystals proved to be organic calcium salts, probably calcium oxalate. Silicon was associated with the crystal sand containing cells of the mesocarp, either as very small silica grains in the cell or randomly distributed in the cell wall as in normal cells. The long raphid crystals may perforate neighbouring parenchyma cells during picking and transporting of the fruit and hence limit its storage life-time. Nutrient substances used in substrate culture, especially those with a high nitrate-ion concentration, may be partly responsible for the formation of the calcium oxalate crystals.     

The impact of changing light levels and fruit load on the pattern of tomato yields

Summary

Experiments in glasshouse and controlled environment facilities investigated the effect of different fruit removal and lighting/shading treatments on the pattern of tomato yields. While the removal of flowering trusses resulted in a yield loss about eight weeks later, there was little loss in cumulative yield as assimilates were distributed to neighbouring trusses. In the growth room experiment, increased photosynthetic photon flux density (PPFD) for one week resulted in a period of increased yield from 4–6 weeks after the start of the treatment, followed by suppressed yields due to smaller fruits on subsequent trusses. However, neither fruit load nor assimilate availability appeared to be responsible for the fluctuations in yield recorded within the glasshouse crop. In this experiment fruit size remained fairly consistent (except when fruit removal treatments were applied), whereas the number of fruits picked per week exhibited much greater variability. This was the case even when all trusses were pruned to leave five fruits, and so was not due to a cycle in the number of fruits set per truss. The flushes in yield were found to be a consequence of a hastening of fruit maturation.     

Thidiazuron increases current-year fruit size and production in Actinidia deliciosa without decreasing return bloom

Summary

The effects of whole-crown application of thidiazuron (TDZ), applied two weeks after full bloom, at the concentrations of 0 (control), 2 or 10 ppm, on vine yield, fruit quality and on next-season potential yield of Actinidia deliciosa ‘Hayward’ were evaluated in 1998–99. Fruit growth increased rapidly after TDZ application. Fruit fresh weight and yield increased by about 13% and 22% with the 2 and 10 ppm concentration, respectively. These responses were associated with only small changes in fruit shape, with 10 ppm-treated fruits slightly rounder than the control. During ripening, TDZ did not affect fruit soluble solid content, flesh firmness or concentration of non-structural carbohydrates. No effect was observed on the amount of winter pruning, entity of bud burst, percent of fertile buds, number of inflorescences per fertile bud or percent of vegetative buds. These results show that TDZ can be applied to the whole crown to increase fruit size and yield, without affecting fruit quality or the next-season’s crop.     

The Growth of Apple Central Fruits as Affected by Thinning with NAA,BA and Naphthenic Acids

Plant growth regulators such as α?naphthylacetic acid (NAA) or 6?benzyladenine (BA) are commonly used for thinning apple fruits. NAs exhibits an auxin- and gibberellin-type biological activity, stimulate the uptake and translocation of auxins in plant tissue, as well as IAA synthesis and act as surfactants. It was assumed that NAs applied at the stadium of early fruit growth might cause fruit abscission in apple, similarly to NAA. The goal of the research was to test the presumed thinning effect of naphthenic acids on apple cultivars ‘Golden Delicious’ and ‘Red Delicious’. Three treatments were applied on both apple cultivars including 16.5?μL L^?1 NAA, 200?μL L^?1?BA and 2.63?mg L^?1?potassium salts of NAs. Cortical cell size and the number of cell layers were measured in order to determine the contribution of cell size and cell division in early fruit growth, as affected by chemical thinning treatments. The effects of plant growth regulators NAA, BA and NAs on fruit set, fruit size and the anatomy of the fruit cortex highly depend on varietal differences. In ‘Golden Delicious’ fruit set reduction by chemical thinning increases both the number of cell layers and cell size of apple fruit, while in ‘Red Delicious’ NAA inhibits fruit growth, while BA is ineffective. Cell growth promoting activity of NAA and NAs, accompanied by the decrease in fruit set and reduced competition among fruits within the canopy, results in a significant fruit weight increase at harvest. The research confirms the auxinic properties of NAs, and their assumed thinning activity.