Effects of Temperature and Photoperiod on Vegetative and Reproductive Growth of Groundnut (Arachis hypogaea L.)*

Effects of temperature × photoperiod interaction on vegetative and reproductive growth were examined in three selected groundnut genotypes by growing them in controlled-environment growth chambers with three temperature regimes (22/18, 26/22, and 30/26°C, day/night) under long (12 h, long day), and short (9 h, short day) photoperiods. The effect of photoperiod on the total dry-matter production (TDM) was significant with the genotypes producing 32–72% greater dry matter under LD than SD. Temperature × genotype interaction effects were significant, with the dry-matter production being greatest at 26/22°C and least at 30/26°C and 22/18°C in two of the three genotypes. Leaf area (LA) was greater under LD than SD at all temperature regimes. LA accounted for 76% of the variation in shoot + root dry weight (R²= 0.76, P < 0.01). A lack of relationship between LA and pod weight or pod numbers suggested that the pod development is controlled by factors other than carbon assimilation. The temperature × photoperiod interaction was significant for root growth, with the root weight being maximal and photoperiod effects being minimal at 22/18°C, while at 26/22°C, root weight declined and photoperiod effects became prominent. Low temperature (22/18°C) affected the reproductive development by reducing the proportion of reproductive nodes in total (vegetative + reproductive) nodes. The conversion of pegs into pods, as indicated by pod to peg ratio (PPR), was lower in LD than in SD conditions. Results suggested that the PPR could be used as an indicator of genotypic sensitivity to photoperiod in groundnut.     

Low-temperature-related growth and photosynthetic performance of alloplasmic tomato (Lycopersicon esculentum Mill.) with chloroplasts from L. hirsutum Humb. & Bonpl.

Growth and photosynthetic performance were analyzed in alloplasmic tomato at a high- (25/17 ^°C; HTR) and low-temperature regime (12/6 ^°C; LTR) in order to establish the role of cytoplasmic variation on low-temperature tolerance of tomato (Lycopersicon esculentum Mill.). Four alloplasmic tomato lines, containing the nuclear genome of tomato and the plastome of L. hirsutum LA 1777 Humb. & Bonpl., an accession collected at high-altitude in Peru, were reciprocally crossed with 11 tomato entries with a high inbreeding level and a wide genetic variation, resulting in a set of 44 reciprocal crosses. Irrespective of growth temperature, alloplasmic families with alien chloroplasts of L. hirsutum (h) were on average characterized by a high shoot biomass, a large leaf area, and a low specific leaf area in comparison with their euplasmic counterparts. These results do not directly point to an advantageous effect of h-chloroplasts on biomass accumulation at low temperature but rather towards a small general beneficial effect on growth and/or distribution of assimilates. Significant chloroplast-related differences in photosynthetic performance, however, were not detected at both temperature regimes, indicating that h-chloroplasts can properly function in a variable nuclear background of L. esculentum. It is concluded that chloroplast substitution is not an effective method for breeding tomato plants with improved low-temperature tolerance. This revised version was published online in August 2006 with corrections to the Cover Date.     

Vitamin C in broccoli (Brassica oleracea L. var. italica) flower buds as affected by postharvest light,UV-B irradiation and temperature

In this study, the changes in vitamin C, l-ascorbic acid (AA) and l-dehydroascorbic acid (DHA) levels in broccoli flower buds were examined during pre-storage and storage periods, simulating refrigerated transport with wholesale distribution and retail, respectively. Broccoli heads were pre-stored for 4 or 7 days at 0 °C or 4 °C in the dark and then stored for 3 days at 10 °C or 18 °C. During storage the broccoli heads were exposed for 12 h per day to three different levels of visible light (13, 19 or 25 μmol m⁻² s⁻¹) or a combination of visible light (19 μmol m⁻² s⁻¹) and UV-B irradiation (20 kJ m⁻² d⁻¹), or they were stored in the dark. The vitamin C content in broccoli flower buds during storage was significantly affected by pre-storage period and temperature. Higher vitamin C levels in flower buds after storage were observed for broccoli heads pre-stored for 4 days or at 0 °C as compared to those pre-stored for 7 days or at 4 °C. Storage temperature also affected vitamin C in broccoli flower buds, with higher levels observed for broccoli stored at 10 °C than at 18 °C. Hence, vitamin C in broccoli flower buds was demonstrated to decrease together with increasing pre-storage period, pre-storage temperature and storage temperature. AA in broccoli flower buds was influenced mainly by storage temperature and to a minor extent by pre-storage temperature. The DHA level and DHA/AA ratio were stable in flower buds of broccoli pre-stored for 7 days, whereas increasing tendencies for both DHA level and ratio were observed after pre-storage for 4 days. These results indicate a shift in the ascorbate metabolism in broccoli flower buds during storage at low temperatures, with its higher rate observed for broccoli pre-stored for shorter time. There were no effects of the light and UV-B irradiation treatments on vitamin C, AA and DHA levels in broccoli flower buds.     

Expression of thermosensitive genic male sterility in rice under varying temperature situations

Response of thermosensitive genic male sterility in rice to varying temperature situations was studied by using four TGMS lines. In three sets of maximum, minimum and their combined temperatures, it was observed that maximum temperature played a predominant role in influencing sterility/fertility of TGMS lines under the combined regimes. Expression of a TGMS gene was found to be influenced by the genetic background of the recipient lines. Exposure for more than 8 hours of 32 ^°C was found necessary to induce complete male sterility in indica TGMS lines: IR68945-4-33-4-14 and IR68949-11-5-31 while, more than 4 hours of such exposure was enough to induce sterility in case of their japonica donor Norin PL 12. Sudden interruption with 27 ^°C even for 2 hours under the sterility inducing regime of 32/24 ^°C could induce partial fertility in the line IR68945-4-33-4-14. However, the line ID 24 remained completely sterile even with 10 hours of interruption with 27 ^°C. An interruption with lower temperature of 27 ^°C for 4 hours for even one day induced partial fertility in IR68945-4-33-4-14. The period of four to eight days after panicle initiation was the stage most sensitive to temperature variations. Hybrid rice breeders need to develop numerous genetically diverse TGMS lines, which possess critical sterility inducing temperature of 28 ^°C and are not affected by sudden interruptions with a lower temperature for few hours daily for a couple of days. This revised version was published online in August 2006 with corrections to the Cover Date.     

Cryopreservation of Delphinium pollen at –30 °C

Pollen of garden varieties and species of Delphinium that was cryopreserved at –30 ^°C after 3 hours of air drying and storage on silica gel, had high germination and pollen tube growth in vitroeven after 180-day storage. On the other hand, pollen stored at 25 ^°C showed a marked decrease in the germination rate within 10 days. The best in vitro germination of Delphinium pollen was on a 1% water agar containing 15% sucrose and 0.005 to 0.01% boric acid and at 15 to 20 ^°C. Field pollination with the cryopreserved pollen showed higher fruit and seed set than pollination with pollen stored at 25 ^°C, and was not significantly different from fresh pollen. This revised version was published online in August 2006 with corrections to the Cover Date.     

Interspecific cross of Brassica oleracea var. alboglabra and B. napus : effects of growth condition and silique age on the efficiency of hybrid production, and inheritance of erucic acid in the self-pollinated backcross generation

Interspecific hybrids were produced from reciprocal crosses between Brassica napus (2n = 38, AACC) and B. oleracea var. alboglabra (2n = 18, CC) to introgress the zero-erucic acid alleles from B. napus into B. oleracea. The ovule culture embryo rescue technique was applied for production of F₁ plants. The effects of silique age, as measured by days after pollination (DAP), and growth condition (temperature) on the efficiency of this technique was investigated. The greatest numbers of hybrids per pollination were produced under 20°/15°C (day/night) at 16 DAP for B. oleracea (♀) × B. napus crosses, while under 15°/10°C at 14 DAP for B. napus (♀) × B. oleracea crosses. Application of the ovule culture technique also increased the efficiency of BC₁ (F₁ × B. oleracea) hybrid production by 10-fold over in vivo seed set. The segregation of erucic acid alleles in the self-pollinated backcross generation, i.e. in BC₁S₁ seeds, revealed that the gametes of the F₁ and BC₁ plants carrying a greater number of A-genome chromosomes were more viable. This resulted in a significantly greater number of intermediate and a smaller number of high-erucic acid BC₁S₁ seeds.     

Photoperiod sensitivity gene controlling heading date in rice cultivars in the northernmost region of Japan

Rice is grown in diverse environments at latitudes ranging from 53^°N to 40^°S. In Japan, Hokkaido is the northernmost rice cultivation region(42–45^°N latitude). Only extremely early maturing (heading) varieties that have extremely low photoperiod sensitivity are adapted to this area. Heading date is the most important trait in adaptation to this particular environment. Also, the efficient manipulation of heading date is a crucial component of rice improvement. To determine the genetic basis for heading date among cultivars grown in Hokkaido, the heading behaviour was analyzed. Clear segregations were observed. To estimate the photoperiod sensitivity of the genes involved, the cultivars and F₁ plants from crosses between the cultivars were grown under different daylength conditions. The results indicated that the genes controlling heading date are photoperiod sensitive, suggesting they play important roles in the northernmost rice cultivation regions in Japan, to which only cultivars with extremely low photoperiod sensitivity are adapted. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effect of incubation temperature regimes and culture medium on broccoli microspore culture embryogenesis

Conditions for reliable induction of embryogenesis from isolated microspores were studied in ten genotypes of broccoli. Embryo yields were significantly increased in almost all of the broccoli genotypes by the incubation at 32.5 ^°C for 1 day, than when the standard incubation at 30 ^°C for 2 days was used. Treatments of 48 hours at 32.5 ^°C produced less than optimal results suggesting that broccoli microspores are more sensitive to high temperatures than those of B. napus. The use of the ¹/₂ NLN-13 medium yielded greater number of embryos than the standard NLN-13. The magnitude of the response to the redution of the concentration of major salts by half in the NLN medium varied with the different genotypes. High embryogenic broccoli cultivars, such as ‘Shogun’, ‘SDB9’, and ‘Green Valiant’, presented a better response to the reduction of the concentration of major salts by half in NLN-13. Reduction never produced a detrimental effect on embryo yield and seems not to have any effect in the subsequent development of embryos in plants. This revised version was published online in July 2006 with corrections to the Cover Date.     

A stability analysis of time to flowering as a screen for responsiveness to temperature and photoperiod in cowpea (Vigna unguiculata)

Summary Twenty-one genotypes of cowpea (Vigna unguiculata), comprising landraces and varieties, were grown in 22 photothermal environments in Nigeria and Niger, West Africa, and a stability analysis of days from sowing to flowering (f) was carried out. Cowpeas are rarely insensitive to photoperiod; they are typically quantitative shortday plants wherein f is delayed when photoperiod (P) is longer than the critical photoperiod (P _c ). Therefore, in order to quantify genotypic variation in temperature sensitivity, genotype f was regressed against the mean trial f in circumstances where P c (i.e. approximately 13 hd^-1) and mean temperature (T) was between 19° and 28° C. Correspondingly, in order to assess genotypic variation in photoperiod sensitivity, trials where T was near optimal (25°–28° C) but where P ranged from 10–14.5 hd^-1 were used. These stability analyses detected no significant differences (P>0.05) between genotypes 9n temperature sensitivity but revealed significant differences (P<0.001) in photoperiod sensitivity. Regression coefficients from the stability analysis were strongly correlated (r=0.94, 19df) with a photoperiod sensitivity constant, c, determined from a photothermal flowering model. A stability analysis of f from field trials can therefore identify and quantify genotypic variation in response to temperature and photoperiod in cowpea.Abbreviations f days from sowing to flowering - P mean photoperiod - P _c critical photoperiod - P _ce ceiling photoperiod - T mean temperature - T _b base temperature - T _o optimum temperature - SDP short-day plant     

In vitro selection for oleic and linoleic acid content in segregating populations of microspore derived embryos of Brassica napus

Microspore derived embryos (MDEs) in Brassica napuscontain large amounts of storage lipids which show a genotype specific fatty acid composition (FAC). One cotyledon of regenerating emblyos can be dissected at an early stage during the in vitro culture and used for fatty acid analysis. Thus, in breeding programmes to modify oil quality, only MDEs having the desired FAC need to be regenerated to plantlets and transferred to the greenhouse. In the present study the applicability of this method for the selection of a high oleic acid content and a low linoleic acid content in the seed oil has been tested by crossing a Brassica napus mutant line having a high oleic acid (C18:1) content in the seed oil (75%) with a wild type doubled haploid line with 62% C18:1 in the seed oil. Microspore culture was applied to the F1 plants. In total 59 MDEs were obtained, from which 31 were cultured with and 28 without 15μM abscisic acid for 3 weeksin vitro. One cotyledon was dissected under aspetic conditions and used for fatty acid analysis. The remaining part of the embryos were further regenerated to plantlets and transferred to the greenhouse to obtain seeds after self pollination. Seeds harvested from the doubled haploid lines in the greenhouse were used for fatty acid analysis and also for growing in the field. The abscisic acid treatment of the MDEs generally improved the correlations for linoleic and oleic acid between the MDEs and the seeds harvested in the greenhouse and the field. The correlations ranged from 0.68** to 0.81**.This indicates that selection for high oleic acid can be started already during an early stage of the in vitro culture. This revised version was published online in July 2006 with corrections to the Cover Date.     

Factors affecting haploid production in wheat using the Hordeum bulbosum system. 3. Post-fertilization effects on embryo survival

Summary An investigation was made of factors affecting the frequency of development of fertilized ovules following wheat x Hordeum bulbosum hybridizations. Seed survival, a term used here to represent the relationship between seed set and fertilization, was shown to vary between the wheat genotypes tested, but was unrelated to the crossability genes present within the wheat genotype. A high seed survival was obtained in crosses involving the variety TH3929 and the single chromosome substitution line, Chinese Spring (Hope 5B). Chinese Spring showed a poor seed survival. Some influence of the H. bulbosum genotype on seed survival was detected. The two environmental factors studied, namely the post-pollination application of gibberellic acid and the ambient temperature, were both shown to affect seed survival. The application of gibberellic acid immediately after pollination or 1 and 2 days after pollination improved seed survival, although the extent of the response was dependent upon the number and timing of the application(s). A genotype-dependent response to the ambient temperature was observed, TH3929 showing a slight but insignificant reduction and Chinese Spring a significant improvement in seed survival with an increase in temperature from 20°C to 26°C. The physiological processes which may be influenced by GA and the ambient temperature and may thereby affect seed survival are discussed.     

Environmental effects on genetic variation of chilling resistance in cucumber

Environmental effects on genetic variation for chilling resistance were studied in nine cultivars and breeding lines (referred to as cultigens hereafter) of cucumber (Cucumis sativus L.). Five experiments were carried out in controlled-environment chambers to measure the effects of growth temperature, photoperiod, duration of chilling, light level during chilling, and watering frequency on chilling resistance of seedlings at the cotyledon and first true leaf growth stages. Significant interactions were found between cultigen and all environmental factors studied except for the photoperiod and watering frequency. Cultigen rank was affected by growth temperature before chilling, chilling duration, and light level during chilling, but shifts in rank were not consistent. Genetic variation was largest when the plants were grown at 22/18 °C, most pronounced after a chilling duration of 5 to 9 hours and a light level during chilling of 270 μmol·m^-2·s^-1. Variation was larger at the first true leaf stage than at the cotyledon stage. Differences among cultigens in chilling damage were largest 5 days after chilling. Therefore, it seems that testing for genetic variation in chilling damage can be restricted to one set of environmental conditions. We recommend the following conditions for screening cucumber for genetic variation in chilling resistance: grow the plants at 22/18 °C, under a 9-hour photoperiod with a 3-hour night interruption, water them once daily, subject them at the first true leaf stage to a chilling treatment of 7 hours at 4°C at a light level of 270 μmol·^-2·s^-1, and evaluate damage 5 days after treatment. This revised version was published online in July 2006 with corrections to the Cover Date.     

A possible gene source for early,day-length neutral pigeon peas,Cajanus cajan (L.) Millsp.

Summary Atylosia scarabaeoides Benth. and A. platycarpa Benth., close relatives of the species Cajanus cajan (L.) Millsp., were screened for photoperiodic response. Four photoperiods ranging from 12 h 45 min to 19 h were studied in three environments. A. scarabaeoides flowered freely only in the first photoperiod. A. platycarpa not only flowered early (39 to 63 days after planting) in all four photoperiods, but also exhibited a relatively constant vegetative phase up to 16 h of illumination. Cajanus cajan, in which most cultivars-if not all-exhibit a response to photoperiod, was successfully hybridized with the two Atylosia species. It is suggested that genes for earliness and insensitivity to day-length could be transferred from A. platycarpa to C. cajan.     

Introgression of unique characteristics of floral initiation under 24 hour day-length of Fragaria chiloensis ‘CHI-24-1’ into F. × ananassa

A unique flower initiation of the wild Junebearing strawberry strain ‘CHI-24-1’ in Fragaria chiloensis occurs under 24 h DL and high temperature conditions. To introduce the floral initiation characteristics of ‘CHI-24-1’ into cultivated strawberries of F. × ananassa, a cross pollination was conducted between ‘CHI-24-1’ and the Japanese short-day type strawberry cultivar ‘Nyoho’. The floral initiation of ‘CHI-24-1’ was induced in both parent and daughter plants linked with runners under a 24 h DL and 23/20 ^∘C, but not 8 and 16 h DLs at the same temperatures. Of the 21 F1 hybrids grown under the 24 h DL, 12 showed flower truss production in the parent and/or daughter plants linked with runners. Among 64 F1 hybrids, 26 exposed to 8, 16 and 24 h DLs for 30 days produced no flower trusses. However, 32, one and five F1 hybrids produced flower trusses under the 24 h DL alone, 8 h DL alone and both the 8 and 24 h DLs, respectively. The results of the experiments indicated that none of the F1 hybrids were day-neutral plants, but approximately 60% had the characteristics of floral initiation under 24 h DL, which was inherited from the pollen parent of ‘CHI-24-1’. The importance of the unique floral initiation characteristics under 24 h DL with high temperature for strawberry breeding was discussed.     

Inheritance of temperature sensitivity of the photoperiod response in common bean (Phaseolus vulgaris L.)

Summary Photoperiod response of flowering in common bean (Phaseolus vulgaris L.) is thought to be controlled by the genes Ppd and Hr. However, cultivars also vary in the degree that cooler temperatures reduces their sensitivity to photoperiod. To examine the inheritance of this temperature sensitivity, crosses of cvs. Gordo x de Celaya and Flor de Mayo × Rojo 70 were evaluated at two sites differing in mean temperature and using 12.5-h natural photoperiod or 18-h artificially extended photoperiod. Under 18-h photoperiod at the warmer site, Palmira, no plants of the parents or of the F₂ populations flowered, confirming that the parents were sensitive to photoperiod. Under 12.5-h photoperiod at the cooler site, Popayan, the parents for each cross flowered at similar dates and no segregation for days to flower was observed. However, under 18-h photoperiod, de Celaya and Rojo 70 and the F₁ populations did not flower within 100 days after planting, while the F₂ and F₃ populations showed segregation that was consistent with single gene inheritance, late flowering being dominant. Late flowering at Popayan under 18-h photoperiod indicates a lack of temperature sensitivity, so temperature insensitivity of the photoperiod response was dominant to sensitivity. The name Tip, for temperature insensitivity of photoperiod response, is proposed for this gene, with the recessive form of this gene conditioning earlier flowering at cooler temperatures with long daylengths. It is recognized that the observed segregation patterns could represent the effect of multiple alleles at the Ppd or Hr loci, and studies are proposed to test this possibility with molecular markers and recombinant inbred lines.     

In vitro screening of strawberry plants for cold resistance

Formation of embryo autonomy of strawberry, plant regeneration fro membryo components, plant freezing conditions in vitro and the possibility to differentiate objectively genotypes by freezing them in vitro and in vivo were studied to create strawberry screening technology in vitro for cold resistance. It was established that autonomy of strawberry embryos manifests itself not earlier than on 14–16^th day after pollination and full autonomy is reached on 20–22^nd day. Plants regenerated from 26 days old embryos grew most intensively. At the highest rate strawberry plants regenerated from an isolated embryo axis on MS medium without phytohormones, and from rescued cotyledons x on the medium with 1.0 BA and 0.5 NAA. The temperature interval, at which genotypes differentiated according to cold resistance in vitro, was -8 to 12 ^°C. Differentiation of strawberry genotypes according to this character conformed to their differentiation in vivo, provided hardening proceeded not less than 21 days. The correlation between cold resistance in vitro and in vivo reached 0.93. Domination of cold resistance manifested itself in strawberry seedlings from various crossing combinations. This revised version was published online in August 2006 with corrections to the Cover Date.     

Slow-growth conservation of potato microplants: efficacy of ancymidol for long-term storage in vitro

Ancymidol was investigated as an alternative mediumsupplement to mannitol for slow-growth conservation ofpotato microplants in vitro. Differentconcentrations of ancymidol (0, 5, 10, 15, 20, 25, 30,35 and 40 M) were tested in slow-growthmedia based on MS medium supplemented with either 30or 60 gl^-1 sucrose. The cultures were conservedunder a 16-h photoperiod at two temperature regimesi.e. 24 ± 1 ^°C and 6 ± 1 ^°C. Therewere significant interactions between ancymidol andother factors such as sucrose, temperature andgenotype for microplant survival, microshoot heightand overall microplant growth. Ancymidol did have abeneficial effect on culture viability after prolongedmaintenance in vitro. The growth-inhibitingeffect of ancymidol persisted through a 16-monthculture period. Combined effect of ancymidol, sucroseand temperature showed that optimum culture viabilityand desirable microplant growth were obtained when thecultures were grown in MS medium supplemented with 10M ancymidol plus 60 gl^-1 sucrose at6 ± 1 ^°C. Vitrification and flaccidity, whichare very frequently observed in potato microplantcultures during prolonged maintenance in vitrounder osmotic stress (mannitol), were not observedwhen the microplants were conserved in ancymidolmedia. Genetic stability of potato microplantsconserved in ancymidol media was evaluated usingrandomly amplified polymorphic DNA (RAPD)fingerprints. Ancymidol did not induce any detectablegenetic variation in genomic DNA as visualized by theabsence of either any additional RAPD fragment oralterations in RAPD fragment patterns.     

Search for resistance to gummy stem blight (Didymella bryoniae) in cucumber (Cucumis sativus L.)

Summary Resistance to gummy stem blight (Didymella bryoniae) was not detected among 1208 cucumber lines of diverse origin when cotyledons of four-day-old seedlings were inoculated with an aqueous suspension of 5000 spores of D. bryoniae delivered to freshly crushed tissue, followed by 48 h incubation at 20°C and 100% RH in the dark. Among 49 lots evaluated for gummy stem blight in the field, cv. Homegreen #2 and Pl 200818 from Burma were resistant. Earliness in fruit maturity was positively correlated with susceptibility to gummy stem blight (r=0.424). Cucumber accessions differed in the incidence of stem and hypocotyl cankers. In greenhouse grown plants at the 2nd–3rd leaf stage, differentiation of resistant lines was obtained by spraying plants with an aqueous suspension of 10⁶/ml D. bryoniae spores followed by 48 h incubation at 20°C and 100% RH in the dark. One cycle of mass selection in Homegreen #2 resulted in improved gummy stem blight resistance.     

Factors affecting in vitro flowering and fruiting of green pea (Pisum sativum L.)

Multiple shoots were efficiently regenerated from cotyledonary node and shoot tip explants of Pisum sativum within 15 days on MS medium containing B₅ vitamins and supplelmented with 2.0 mgl^-1 6-benzylaminopurine. The elongated shoots produced on the same medium were excised and transferred to MS medium containing half strength ammonium nitrate (8.25 gml^-1) and supplemented with auxins (indole-3-butyric acid or naphthalene acetic acid) either alone or in combinations with gibberellic acid. Rooting and flowering were observed on the 7th and 15th day after their transfer to rooting medium. The flowers self-fertilised in vitro and produced mature pods within 25 days of rooting. These seeds were germinable both in vitro and in vivo. In vitro seeds sown in pots under field conditions developed into flowering plants, and subsequently produced pods with viable seeds. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of net photosynthesis,dark respiration,stomatal resistance and related characters in tomato (Lycopersicon esculentum Mill.) under low energy conditions

Summary Inheritance of net photosynthesis, dark respiration, specific leaf fresh weight, and stomatal resistance was studied in a diallel cross without reciprocals with 15 tomato genotypes. The plants were grown under low light intensity (24 Wm^-2), short days (8 h) and low temperature (19 °C/10 °C day/night temperature). Significant f-values for GCA were observed for all characters. Smaller, but for most characters also significant f-values for SCA were found. This indicated that the characters were mainly controlled by additive gene action. The relationships between the physiological characters and breeding aspects are discussed.