Effects of leaf aging and light duration on photosynthetic characteristics in a cucumber canopy

Photosynthetic characteristics, chlorophyll index and leaf area were examined in selected leaves of cucumber (Cucumis sativus L. cv. Euphorbia). In the first experiment, plants of cucumber were grown horizontally at a lighting period of 20 h day⁻¹. Photosynthetic measurements in horizontally growing cucumbers showed that there was no decline in photosynthetic capacity when cucumber leaves are developing under good light conditions. In a second experiment, plants were grown in a traditional high-wire cultivation system under 20 h day⁻¹ lighting period until they reached final height and then exposed to different lighting periods (20 and 24 h day⁻¹) for 3 weeks. In stands of cucumber plants photosynthetic measurements showed that the lower leaves have a significant reduction in photosynthetic capacity due to reduced light conditions. Three weeks exposure to 24 h day⁻¹ lighting period reduced leaf area by 20%. Plant grown under continuous light had also lower chlorophyll index compared to plants grown under 20 h day⁻¹ lighting period.     

Two potent allelopathic substances in cucumber plants

Since cucumber plants are mostly discarded as large waste after crop harvesting, allelopathy of cucumber plants was investigated for possible weed management options and utilization of the waste. Two potent growth inhibitory substances were isolated from an aqueous methanol extract of cucumber (Cucumis sativus L. cv. Phung Tuong) plants. These substances were determined as 9-hydroxy-4,7-megastigmadien-9-one (HMO) and (6S,7E,9S)-6,9,10-trihydroxy-4,7-megastigmadien-3-one (THMO) by the analysis of MS, ¹H NMR spectra and optical rotation. HMO inhibited the growth of cress (Lepidium sativum L.) and Echinochloa crus-galli (L.) Beauv seedlings at concentrations greater than 0.3 and 1 μM, respectively. THMO inhibited the growth of cress and E. crus-galli seedlings at concentrations greater than 1 and 3 μM, respectively. The concentrations required for 50% growth inhibition on roots and shoots of cress and E. crus-galli were 2.4–29.3 μM for HMO and 8.1–52.2 μM for THMO. The endogenous levels of HMO and THMO in cucumber plants were 31.8 and 43.5 μg g⁻¹ dry weight, respectively. These results suggest that HMO and THMO may be the causal factors for the growth inhibitory effect of cucumber plants. Therefore, cucumber plants may be potentially useful for weed management options in an agricultural setting, such as a cover crop and soil admixture, which should be investigated further in the field.     

Effects of interlighting on yield and external fruit quality in year-round cultivated cucumber

The effects of interlighting and of the proportion of interlight on the yield and fruit quality of year-round cultivated cucumber (Cucumis sativus L. cv. Cumuli) were investigated for this study. Artificial lighting was provided by high pressure sodium (HPS) lamps and the lighting regimes included top lighting (TL), top + interlighting 24% (T + IL24) and top + interlighting 48% (T + IL48). In TL, all of the lamps were mounted above the canopy. In T + IL24 and T + IL48, top lamps covered 76 and 52% of the lighting, respectively, while 24 and 48% of the lighting came from interlighting lamps which were mounted vertically 1.3 m above the ground between the single plant rows. The outdoor daily light integral (DLI) varied greatly during the cultivation periods; the mean values were 36.8, 5.3 and 19.9 mol m⁻² day⁻¹ for the summer, autumn–winter and spring stands, respectively. Lighting regime affected both yield and external fruit quality. Interlighting increased first class yield and decreased unmarketable yield, both in weight and number. The increase in the annual first class yield in weight was 15% in the two T + IL regimes. Interlighting improved energy use efficiency in lighting, being for the whole year 120, 130 and 127 g total yield kW h⁻¹ in TL, T + IL24 and T + IL48, respectively. Interlighting increased the fruit skin chlorophyll concentration in all seasons, but had only a minor effect on the fruit dry matter concentration. The mean total chlorophyll concentration in fruit skin was 70.8, 76.7 and 82.2 μg cm⁻² in TL, T + IL24 and T + IL48, respectively. In addition, interlighting extended the post-harvest shelf life of cucumber fruits in spring. Besides interlighting per se, also the higher proportion of interlight tended to further improve the fruit quality. For example, the fruit skin chlorophyll concentration increased along with the higher proportion of interlighting. In general, the effects of lighting regime were more prominent in lower natural light conditions in winter and spring. It is concluded that interlighting is a recommendable lighting method in cucumber cultivation, especially in lower natural light conditions.     

Effects of intracanopy lighting on photosynthetic characteristics in cucumber

Plants of cucumber (Cucumis sativus L. cv. Euphorbia) were grown in a traditional high-wire cultivation system to investigate the effects of three lighting regimes on photosynthetic characteristics, leaf area and yield. The lighting regimes included overhead lighting (OH), where all the lamps were mounted above the canopy and overhead + intracanopy lighting (OH + IC) which comprised 65% of overhead lamps and 35% of lamps mounted vertically along the plant rows. All overhead lighting was provided for 20 h day⁻¹ and intracanopy lighting was provided for either 20 h or 24 h day⁻¹ lighting period. Intracanopy lighting improved the light distribution in the canopy. Gas exchange measurements showed that intracanopy lighting increased net photosynthesis (P_N) and photosynthetic capacity (P_max). Parameters calculated from CO₂ response (A/C_i) curves showed that in vivo estimate of the maximum rate of Rubisco carboxylation (Vc_max) and the maximum rate of electron transport (J_max) were affected by light regime. Intracanopy light increased yield by 11% compared to traditional overhead light.     

High frequency shoot regeneration from cotyledon explants of cucumber via organogenesis

Organogenic callus induction and high frequency shoot regeneration were achieved from cotyledon explants of cucumber. About 86.2% of cotyledon explants derived from 5-day-old in vitro raised seedlings produced green, compact nodular organogenic callus in MS medium containing NAA (2.69 μM) and BA (4.44 μM) after two successive transfers at 20 days interval. Adventitious shoots were produced from the organogenic callus when it was transferred to MS medium supplemented with NAA (1.34 μM), BA (8.88 μM), zeatin (0.91 μM) and l-glutamine (136.85 μM) with shoot induction frequency of 75.6%. Shoot proliferation occurred when callus with emerging shoots was transferred in the same medium at an interval of 20 days. Shoots (1.0 cm length) were excised from callus and were elongated in MS medium fortified with GA₃ (1.44 μM) and BA (4.44 μM). The elongated shoots were rooted in MS medium supplemented with IBA (3.42 μM) and BA (4.44 μM). Rooted plants were acclimatized in green-house and subsequently established in soil with a survival rate of 80%. This protocol yielded an average of 35 shoots per cotyledon explant in a culture duration of 120–140 days.     

Fertilization regulates soil enzymatic activity and fertility dynamics in a cucumber field

Different fertilizers may affect soil enzymatic activity and soil fertility dynamics. These effects were investigated in a field experiment with cucumber (Cucumis sativus L.) and the relationship with yield and soil nutrient availability was assessed. Soil enzymatic activity, measured as phosphatase, catalase, invertase and urease activities, decreased in the early growth stages of cucumber, but increased in the late ones, when plant were supplied with partially decomposed horse manure. Chemical N fertilizer inhibited soil enzymatic activity but P and K fertilizers enhanced it. Activity of different soil enzymes was positively correlated with soil NH₄⁺–N and available P concentration, but negatively correlated with leaf N and P concentration. Cucumber yield was also positively correlated with the soil enzymatic activity. Our results demonstrate that soil enzymatic activity acted as a useful indicator of soil fertility dynamics.     

Measurement system of whole-canopy carbon dioxide exchange rates in grafted cucumber transplants in which scions were exposed to different water regimes using a semi-open multi-chamber

A continuous CO₂ measurement system was developed to monitor the CO₂ exchange rate of the whole canopy of grafted transplants using semi-open multiple chambers. Air heating or cooling and humidification inside a healing box were under control, if needed. To test the system, the gas exchange rate of the cucumber (Cucumis sativus L.) transplants grafted onto pumpkin (Cucurbita maxima cv. ‘New-Shintozwa’) was analysed. During the healing and acclimatisation of the grafted cucumber plants, the air temperature in the box remained constant at night but ranged above 1 °C of a set value under high humidity in daytime. The relative humidity was kept within the set point during the daytime, and it nearly reached 100% at night when not controlled. The cucumber seedlings were exposed to different water stresses before grafting, and the water potentials of each treatment were −0.579 (non-stressed), −0.814 (mildly water-stressed), and −0.870 MPa (strongly water-stressed) on grafting. At the water-stressed scions, leaf expansion was inhibited by 30.9–53.3% compared with the non-stressed scions. Therefore, the gas exchange rates of the strongly water-stressed scions based on the leaf area were decreased to 72.7% compared with the non-stressed scions. After grafting, the apparent photosynthesis of the transplants of all treatments was negative, with higher respiration in the strongly water-stressed scions during the initial period of healing. However, they turned to positive values and exceeded those of the non-stressed scions from three days after grafting. This result provides critical information that the water column is physiologically connected between the stock and scion within two days after grafting. As a result of water stress, the leaf area and dry weight of the transplants in the strongly water-stressed scions were inhibited by 67.5% and 83% compared with the non-stressed scions at the end of acclimatisation. In contrast, the relative growth rate and graft-take of the strongly water-stressed transplants were slightly increased, which suggests that the water stress prior to grafting alleviated the water demand of the scion. This system may provide useful information for diagnosis at the early stage by monitoring the whole canopy's photosynthesis over a long term.     

Efficient embryo induction in cucumber ovary culture and homozygous identification of the regenetants using SSR markers

Several factors, i.e. the duration of thermal shock pretreatment at 35 °C, the concentrations of TDZ and the silver nitrate were investigated for their effects on embryo formation in a variety of cucumber (Cucumis sativus L) ovary culture. The results showed that a thermal shock for 3 days at 35 °C at the start of the culture resulted in higher frequency of embryo formation than 2 or 4 days. TDZ had a positive effect on the embryo formation. Highest embryo formation frequency (72.7%) was recorded by adding 0.04 mg/L TDZ into the induction medium. The results found that addition of AgNO₃ to induction medium had no significant effect on frequency of embryo formation but shortened embryo sprouting period and improved number of embryos formed in each ovary slice. All the experiment materials responded well to ovary culture, and there were no difference among genotypes used in this study. Among the forty regenerated plants obtained, two were identified as haploid plants (2n = x = 7), five were tetraploid plants (2n = 4x = 28), and the rest were diploid plants. Microsatellite markers (SSR) were used to analyze the homozygosity of the diploid plants, the putative chromosome-doubled haploids. Of the 33 diploid plants, 17 (51.5%) were identified as double haploids. Based on the above results, we have established a useful protocol for production of cucumber doubled haploids with ovary culture.     

The inheritance of two novel subgynoecious genes in cucumber (Cucumis sativus L.)

Cucumber (Cucumis sativus L.), which is a vegetable crop, has served as the model system for sex expression in flowering plants, and the inheritance of sex expression in cucumber is well documented. However, the genetics of subgynoecism expression in cucumber had rarely been described. In this study, we investigated the inheritance of subgynoecious traits in cucumber plants with the inbred cucumber lines of subgynoecious (C. sativus L. var sativus cv 97-17 and S-2-98) as the materials. Genetic analysis had showed the two subgynoecious inbred lines were controlled by one pair of recessive gene and one pair of incompletely dominant gene, which were designated presently as mod-F2 and Mod-F1, respectively. Furthermore, the mod-F2 and Mod-F1 loci, which enhance the intensity of femaleness, also inherited independently with F and M genes.     

Salicylic acid regulates sugar metabolism that confers tolerance to salinity stress in cucumber seedlings

Salicylic acid (SA) plays a critical role in plant development and defense responses to biotic and abiotic stresses. Sugars can act as osmotic adjustors or nutrient and metabolic signals in the activation of plant defense responses. To uncover the effects of SA on sugar metabolism, we assayed the altered components and levels of sugars in cucumber seedlings treated with SA. After SA treatment, the levels of glucose, fructose, raffinose and stachyose were increased in both leaves and roots. In contrast, sucrose and starch responded differently in leaves and root, decreasing in leaves but accumulating in roots. These changes could be due to the effects of SA on the activities of metabolism-related enzymes. In leaves, SA promoted the activities of sucrose phosphate synthase (SPS), sucrose synthase (SS) and amylases, while the SA-treated root showed a reduced amylase activity and an unchanged activity in SPS. The changes in various sugar contents resulted in the accumulation of soluble sugars in SA-treated cucumber seedlings, especially non-reducing sugars in roots. These increased sugars could function as osmotic regulators and facilitate water uptake and retention in plant cells, thereby conferring seedlings an enhanced tolerance to salinity stresses caused by NaCl treatment. Taken together, our findings provide an important insight into the effects of SA on sugar metabolism, and a protective mechanism for SA against water deficiency is discussed.     

Responses of cucumber plant to NH4 and NO3 nutrition: The relative addition rate technique vs. cultivation at constant nitrogen concentration

Different N sources (NO₃⁻, NH₄⁺, or NH₄NO₃) at different relative addition rates (RAR) were supplied to cucumber (Cucumis sativus L.), a species sensitive to NH₄⁺ toxicity. For comparison, cucumber plants were also grown at constant concentrations of 1 and 5 mM NH₄⁺ or 5 mM NO₃⁻. The fresh weight of NH₄⁺-fed plants at RAR 0.15 and RAR 0.25 day⁻¹ was similar to that of NO₃⁻-fed plants, while at RAR 0.35 or RAR 0.45 day⁻¹ growth reduction occurred. When available as a constant concentration, NH₄⁺ decreased plant growth at 5 mM. It is concluded that at low rates of N supply the relative addition rate technique can be used for growing cucumber plants with NH₄⁺ as sole N source without deleterious effects.     

Plant spacing and cultivar affect melon growth and yield components

The objective of this study was to evaluate the effect of cultivars and in-row spacing on vegetative growth and yield components in melon (Cucumis melo L.). The cultivars ‘Early Dawn’ (ED), ‘Superstar’ (SS) and ‘Medoro’ (ME) were grown in rows 1.5 m apart at plant spacing 0.6, 0.9, 1.2 or 1.5 m. Cultivars ED and SS were grown in 1998 and 1999, while ED and ME were tested in 2000. The main vine length, diameter, number of leaves and lateral branches were highest at 1.5 m in-row spacing in 1999 and 2000, while there was no influence of plant spacing on observed growth parameters in 1998. ED performed better compared with other two varieties, mostly due to more fruit per ha and per plant. Yield was significantly decreased with increased plant spacing in all 3 years of research. At in-row spacing of 1.5 m yield was in average 25% lower than at 0.6 m spacing. Average fruit weight slightly increased with plant spacing up to 1.2 m, even though the differences were significant only in 1999. Small differences among cultivars were also found. Number of fruit per ha decreased linearly while number of fruit per plant increased linearly as plant spacing increased in all cultivars except SS in 1999.     

Increasing cherry tomato yield using fish effluent as irrigation water in Northeast Brazil

The rational use of water in arid and semi-arid zones is fundamental for resource sustainability. The integration of aquaculture with agriculture appears to be an excellent way of saving water, disposing aquaculture effluents and providing additional fertilizer to the agricultural crop. The objective of this study was to test the feasibility of using fish effluent to irrigate cherry tomatoes cultured with different types of organic fertilizers. A field experiment was carried out in a randomized complete block design in a 2 × 5 factorial scheme, with four replications. The treatments consisted of the combination of two irrigation water types (well water and fish effluent) with five fertilizers (cow manure, chicken manure, vermi-compost, commercial compost and a control without fertilization). Evaluations for tomato fruit number, productivity and mean fruit weight were performed. Statistical analyses were done in the following harvest periods: 0–25, 0–50, 0–75 and 0–100% of the harvest days. On the first three harvest periods analyzed, treatments irrigated with fish effluent had higher fruit number and productivity. This effect was more evident when the organic fertilizer used did not attend all plant needs. The higher productivity observed in effluent treatments was related to the increase in fruit number. The higher productivity in treatments with fish effluent has a special importance for small farmers, to whom fertilizers are often cost prohibitive or unavailable.     

Effects of simple and double grafting melon plants on mineral absorption,photosynthesis, biomass and yield

The Spanish type cultivar ‘Piel de Sapo’ (Cucumis melo L. var. saccharinus), has a limited compatibility with the Cucurbita maxima × Cucurbita moschata hybrids currently used as rootstocks. Double grafting can be used to improve compatibility between rootstock and scion by means of an intermediate rootstock compatible with both. Non-grafted, single, and double grafted melon plants of the cultivar Piel de Sapo were evaluated for water, nutrient absorption, photosynthesis activity, biomass production in early phases, as well as for yields and fruit quality in a long term trial. The hybrid ‘Shintoza’ (C. maxima × C. moschata) was used as rootstock, and the cantaloupe type melon cultivar ‘Sienne’ as an intermediate scion. Grafting did not affect net photosynthetic values, yet increased water use efficiency (35%). Double grafted plants increased aerial dry weights (66% and 31% with respect to non-grafted and simple graft plants, respectively), and also increased capacity for uptaking beneficial minerals (between 61% and 13% and particularly for NO₃⁻, P, K, Ca, Mn, and Zn) with respect to non-grafted and single grafted plants. The quantum efficiency PSII photochemistry values increased in double grafted plants (12%) with respect to the control plants. Consequently, double grafting on a vigorous rootstock such as ‘Shintoza’, with an intermediate scion, results in improved mineral and water absorption and achieves an increase in ion influx to the scion – so enabling an increase in light photosynthetic reaction and biomass. Double grafted plants increased fruit yield when compared to simple grafted and non-grafted plants (12% and 56%, respectively) and did not affect fruit quality in terms of °Brix and colour. In conclusion, double grafting presents several beneficial aspects that are counter-balanced by the extra cost of the technique. The difference in yields reflects compatibility problems.     

Physiological responses associated to substrate water availability of Rosa ‘Eurored’ plants grown in soilless greenhouse conditions

The aim of the present study was to provide a better insight into the adaptive processes to water deficit of Rosa hybrid plants grown in soilless cultivation. Rose plants were grown in a greenhouse under two water regimes, corresponding to 100% and 67% of the irrigation needs. Watering was stopped for 0, 72, and 96 h (dry-down period) and photosynthetic parameters, water relations, chlorophyll fluorescence, carbohydrate and ion content were determined.     

Effect of alleviating products with different mode of action on physiology and yield of olive under drought

Two years old self-rooted Koroneiki olive trees (Olea europaea L.) were subjected to two irrigation regimes, i.e. the fully irrigated and the severely water stressed trees, while they were treated with three alleviating products of different mode of action. The products used were the osmolyte glycine betaine, the antioxidant Ambiol and the heat and irradiance reflecting kaolin clay particles. The effects of product application and water regime on leaf characteristics, shoot and root growth, photosynthesis, leaf compatible solids (carbohydrates) concentration and yield were evaluated. All products applied, exhibited significant alleviating action, based on the relative alleviation index. Irrigated trees exhibited greater growth than drought stressed ones, while the ameliorating products maintained the water content of the leaves under drought conditions and resulted in lower leaf tissue density. On the other hand carbon assimilation rate, stomatal conductance and intrinsic water use efficiency were significantly reduced under drought stress, while the opposite stood for intercellular CO₂. Drought stress resulted in elevated sucrose leaf concentration, while the application of Ambiol increased stachyose concentration and that of glycine betaine did the same with the mannitol concentration. Among the alleviating products tested in this experiment Ambiol and glycine betaine had a significant positive effect on leaf water content, photosynthesis and yield under both drought and well irrigated conditions.     

Status of nitrogenous and carbohydrate compounds as affected by nitrogen fertigation rates in young persimmon trees

The effects of nitrogen (N) fertigation rates on levels of nitrogenous and carbohydrate compounds were determined in young ‘Fuyu’ persimmon trees. Total N amounts of 0 g, 10 g, 20 g, 30 g, and 40 g were distributed, respectively, to a 50-L pot by fertigation as different volumes of 1% (w/v) urea solution every week from May 15 to September 25. Fruits were sampled on November 2, and shoots and roots on November 17. High N fertigation rates significantly increased the N and amino acids in all the tree parts and protein in fruits and shoots. In contrast, the concentration of soluble sugars significantly decreased in the fruits and perennial parts as the N rate increased from 10 g to 40 g. A notable reduction of starch was also found in the shoots and roots of trees with high N fertigation rates. The percentage of shoots died due to cold injury during the winter was the highest at the 0 g N rate; however, it was 2.3-fold greater at the 40 g N rate as compared to the 10 g N rate. We concluded that high N fertigation could interfere not only with sugar accumulation of fruits through increased N metabolism, but also reserve carbohydrates of perennial organs.     

Biofertilizers improve plant growth,fruit yield,nutrition, metabolism and rhizosphere enzyme activities of Pomegranate (Punica granatum L.) in Indian Thar Desert

Production of horticultural crops has undergone enormous change in recent years due to development of innovative technologies including integrated nutrient management practices using biofertilizers. The present study represents the positive response of biofertilizers in nursery seedlings followed by their transplantation in harsh field conditions of Indian Thar Desert. Nursery and field experiments were carried out to assess the effectiveness of selected N₂-fixing bacteria and AM fungi alone or in combination, on the growth and biomass production of Punica granatum. In both experiments, the combined treatment of Azotobacter chroococcum and Glomus mosseae was found to be the most effective. Besides enhancing the rhizosphere microbial activity and concentration of various metabolites and nutrients, these bioinoculants helped in better establishment of pomegranate plants under field conditions. A significant improvement in the plant height, plant canopy, pruned material and fruit yield was evident in 5-year-old pomegranate plants in field conditions. In view of the above results, use of biofertilizer technology may be adopted for the establishment and development of other horticultural plant species in arid regions.     

Combined effects of leaf/fruit ratios and N and K fertigation levels on growth and distribution of nutrients in pot-grown persimmon trees

Responses of pot-grown persimmon trees (Diospyros kaki cv. ‘Fuyu’) with two different leaf/fruit (L/F) ratios and three different fertigation levels were studied. On July 2, the L/F ratios of 4-year-old trees were adjusted to 20 and 10 by fruit thinning. From July 4 to August 26, each pot was fertigated with a solution containing 0–0 g (none), 10–8 g (medium), or 20–16 g (high) of nitrogen (N) and potassium (K), respectively. The L/F ratio of 10 resulted in higher yields but smaller fruits containing lower soluble solids than the ratio of 20. The non-fertigated trees with the L/F ratio of 10 produced the smallest fruits, but the medium fertigation increased the size of the fruit at this ratio. The high fertigation level decreased fruit skin color at both ratios. Fruit dry weight increased the most among the tree components, accounting for 48–57% of the tree total dry weight at the L/F ratio of 20 and for 64–72% at the ratio of 10. The increase in tree total dry weight at the ratio of 10 was greater than that at the ratio of 20. The trees receiving medium fertigation had higher tree total dry weights than those receiving high fertigation, especially at the ratio of 10. As the L/F ratio changed to 10 from 20, the percentages of N and K partitioned to fruits in the non-fertigated trees increased from 38% to 51% and from 67% to 96%, respectively, of the tree total N and K increases; the percentage decreased with increasing fertigation level. Although fertigation significantly increased total N and K contents of the tree, the increase in K at the ratio of 10 was less in high fertigation rather than in the medium level. The results indicated that adjusting supplemental N and K was necessary to ensure fruit growth and N and K accumulation for trees with high fruit loads, but high levels of fertigation are not necessarily preferable.     

Nutritional quality of greenhouse lettuce at harvest and after storage in relation to N application and cultivation season

The effect of five levels of nitrogen fertilization on the growth and nutritional quality of Cos lettuce (Lactuca sativa L. cv. Parris Island) at harvest and after storage was studied during autumn and winter in South-West Greece. Plants were cultivated hydroponically in a greenhouse and the nitrate, chlorophyll and ascorbic acid (vitamin C) concentrations within the plant tissues were measured at harvest and following storage at 5 or 10 °C for 10 days. Nitrate accumulated in the leaves with increasing amounts of N within the nutrient solution and was higher in the winter than in the autumn. At the lowest N level (20 mg L⁻¹), the inner leaves accumulated more nitrate than the outer leaves, whereas at higher N levels (140, 200 or 260 mg L⁻¹) nitrate accumulation was higher in the outer leaves. Overall, the highest nitrate concentrations were detected in the petiole and the proximal end of the leaf, but at the lowest N application rate (20 mg L⁻¹) nitrate accumulated in the distal region of the leaf too. Although the nitrate concentrations within the leaves did not change significantly during 10 days storage at 5 or 10 °C, the chlorophyll and vitamin C concentrations decreased. Chlorophyll loss was higher in lettuce that was grown under low N levels and was higher at 10 °C than at 5 °C, but was reduced by enclosure of the lettuce in polyethylene film. It is concluded that the optimum N application rate for Cos lettuce grown hydroponically under cover during autumn and winter in South-West Greece, and in other areas with a similar climate, is 200 mg N L⁻¹ because at this N rate yield is satisfactory and leaf nitrate concentrations are below the maximum acceptable level for human consumption. Nutritional value (vitamin C concentration) and market quality (chlorophyll content) are highest at harvest and decrease during storage, but quality in terms of nitrate concentration does not change.