The inheritance of white fruit and stem color in summer squash Cucurbita pepo L.

Summary White fruited plants from the variety White bush were crossed to green and striped fruit in order to study the inheritance of white fruit in summer squash. The genetic ratio suggested that green fruit is controlled by two genes, C and R, and that one (C) has a dominant epistatic control. The white fruit is determined as ccrr. A third fruit color group appeared in the F₂, F₃ and BC₁, having white fruit after anthesis which changed to green during development. The genetic control for this fruit is suggested to be ccR..The green striped fruit showed a simple dominance (St) over the plain white.The dark stem of the Zucchini variety showed single gene dominance (D) over the light green stem.It was found that an association exists between the nonpersistent fruit color and dark green stem, and between white fruit and light green stem.Contribution from the Volcani Institute of Agricultural Research, Bet Dagan, Israel. 1968 Series; No. 1437E.     

Multiple allelism at a major locus affecting fruit coloration in Cucurbita pepo

The l-2 (light coloration-2) locus in Cucurbita pepo (pumpkin, squash, gourd) has a major effect on fruit color intensity through its complementary interactions with at least three other loci. From a sixth-generation backcross progeny of which a wild gourd having striped fruits was the donor parent and a cultigen having ‘plain light’-colored fruits(genotype l-2/l-2) was the recurrent parent, a true-breeding line was isolated that had a fruit color phenotype that differed from both parents. The fruits of this line were not striped and had a color pattern resembling‘light type 2’ (genotype L-2/–-), but of weaker expression. This newly established line was then crossed with near-isogenic lines of genotypes l-2/l-2 and L-2/L-2 and the resulting filial and backcross progenies were observed under field conditions. The results indicated that the new phenotype is recessive to ‘light type 2’ but dominant to ‘plain light’ and that it is conferred by a single gene. The results of a testcross for allelism indicated that this gene is an allele at the l-2 locus and it is herein designated L-2 ^w. Dominance relationships at the l-2 locus are L-2 > L-2 ^w >l-2. This revised version was published online in July 2006 with corrections to the Cover Date.     

Gene for broad, contiguous dark stripes in cocozelle squash (Cucurbita pepo)

Alternating intense (dark) and light stripes on the fruits are a fairly common feature of Cucurbita pepo, a highly variable species encompassing various kinds of pumpkins, squash, and gourds. The intense stripes can be narrow and broken or broad and contiguous. Narrow, broken intense striping is conferred by allele l-1 ^St, which is dominant to allele l-1 for non-striped, lightly colored fruits and recessive to allele L-1 for non-striped, intensely colored fruits. Cocozelle-type squash often exhibit broad, contiguous intense stripes. The inheritance of this characteristic was studied by transferring it from cocozelle squash and preparing nearly isogenic lines having broad-striped, narrow-striped, and non-striped light-colored fruits. In the broad-striped near-isogenic line, the intense stripes averaged 37% wider than in its narrow-striped counterpart. The broad-striped line was crossed with the narrow-striped line, the non-striped light-colored line, and the non-striped intense-colored `Fordhook Zucchini'. Broad striping was dominant to narrow striping and to non-striped light and conferred by a single gene; it was recessive to non-striped intense. The gene for broad striping was found to be allelic to that for narrow striping and to that for non-stripe dintense and is herein assigned the symbol l-1 ^BSt. Dominance relationships at the l-1 locus were observed to be L-1>l-1 ^BSt >l-1 ^St >l-1. This revised version was published online in July 2006 with corrections to the Cover Date.     

A recessive,hypostatic gene for plain light fruit coloration in Cucurbita pepo

Summary Fordhook Zucchini, a squash (Cucurbita pepo) cultivar having intensely colored fruits conditioned by genotype D/D L-1/L-1 L-1/L-2, was crossed with Beirut, a squash cultivar having lightly colored fruits. Beirut was also crossed with Vegetable Spaghetti, a cultivar having lightly colored fruits conditioned by genotype d/d l-1/l-1 l-2/l-2, and with tester stocks of genotypes d/d L-1/L-1 l-2/l-2 and d/d l-1/l-1 L-2/L-2. The results indicated that Beirut, like Vegetable Spaghetti, possesses genotype d/d l-1/l-1 l-2/l-2. However, no F₂- or backcross-generation segregates possessing the Light type 1 phenotype, conditioned by recombinant genotype d/d L-1/-l-2/l-2 and heretofore observed when Fordhook Zucchini was crossed with Vegetable Spaghetti, were recovered when Fordhook Zucchini was crossed with Beirut. The results of crossing Beirut with Vegetable Spaghetti and with the tester stocks indicated that both Beirut and Fordhook Zucchini carry a recessive gene, designated pl, which results in Plain light fruit coloration in plants of d/d L-1/- l-2/l-2 genotype. Beirut possesses genotype d/d l-1/l-1 l-2/l-2 pl/pl, Fordhook Zucchini D/D L-1/L-1 L-2/L-2 pl/pl, and Vegetable Spaghetti d/d l-1/l-1 l-2/l-2 Pl/Pl. Both the dominant D allele and the dominant L-2 allele are epistatic to gene pl.Contribution No. 3443-E from the Agricultural Research Organization, Bet Dagan, Israel     

Genetic control of irregular striping, a new phenotype in Cucurbita pepo

Longitudinal striping of the fruit exterioris a common trait in Cucurbita pepo(pumpkin, squash, gourd). Striping occursas ten pairs of alternating dark and lightstripes that are regularly situated inaccordance with the ten sub-epidermalcarpellary vein tracts. Usually, the darkstripes occur in the central areas betweenthe vein tracts, with the light stripesoccupying the remainder of the fruitsurface. A new phenotype is described inwhich dark spots or streaks occur in thecentral areas between the vein tracts;these are usually accompanied by darkstreaks or stripes that occur adjacent toand over part or all of the length of someor all of the ten carpellary vein tracts. The inheritance of this irregular stripingpattern was studied by crossing anaccession that bred true for this phenotypewith a near-isogenic line havinglight-colored fruits. The resultsindicated that irregular striping isdominant to light coloration and isconferred by a single gene. Testcrossesfor allelism with regular striping wereprepared by crossing the irregularlystriped accession with near-isogenic lineshaving broad stripes, l-1 ^BSt/l-1 ^BSt, and narrowstripes, l-1 ^St/l-1 ^St. The testcross results revealed thatirregular striping is conferred by anallele at the l-1 locus, designatedl-1 ^iSt. The l-1 ^iStallele is dominant to light-colored (l-1), recessive to dark-colored (L-1), and can be co-dominant or recessiveto broad-striped (l-1 ^BSt) andnarrow-striped (l-1 ^St),depending on whether or not it is fullyexpressed. Dominance relationships at thel-1 locus can, therefore, berepresented as L-1 > (l-1 ^BSt > l-1 ^St) l-1 ^iSt > l-1.     

Shading of plants facilitates selection for powdery mildew resistance in Squash

Summary Powdery mildew development was assessed on squash (Cucurbita pepo) plants of a susceptible cultivar, a resistant accession, their F₁, and their F₂ in an early summer planting in the field, covered or not covered with a shading net. Three reaction types were observed: susceptible, powdery mildew on stems and on both, the upper and lower leaf surfaces, as in the susceptible parent; resistant, no powdery mildew on leaves or stems, as in the resistant parent; and partially resistant, powdery mildew on upper leaf surfaces only, as in the F₁. Disease presence on the stem was associated with susceptibility. Shading hastened the appearance of powdery mildew and increased the severity of infection on partially resistant and susceptible plants, facilitating identification of resistant individuals in the F₂ population.Contribution No. 1613-E from the Agricultural Research Organization, Bet Dagan, Israel     

Postharvest ultraviolet-C irradiation suppressed Psy 1 and Lcy-β expression and altered color phenotype in tomato (Solanum lycopersicum) fruit

Mature green cherry tomato fruit were harvested and treated with ultraviolet-C (UV-C) irradiation at a predetermined dose of 4.2 kJ m⁻², and stored at 18 °C for 35 days. The effects of UV-C treatment on color change, pigment contents, and the expression of major genes involved in carotenoid metabolism, including Psy 1, Pds, Lcy-β, and Lcy-ɛ, encoding phytoene synthase, phytoene desaturase, lycopene β-cyclase and lycopene ɛ-cyclase, respectively, were examined. The UV-C treated fruit developed a pink red color in contrast to the normal orange red color of control fruit. Lycopene accumulation during ripening in UV-C treated fruit was significantly inhibited but its final content was not affected. However, both accumulation and final content of β-carotene were significantly suppressed in UV-C treated fruit. The lower content of β-carotene, leading to a higher lycopene to β-carotene ratio, is probably responsible for the altered color phenotype in UV-C treated fruit. Psy 1, a major gene involved in lycopene synthesis was inhibited by UV-C irradiation. Significantly suppressed expression of Lcy-β gene was also observed in UV-C treated fruit. Thus it is possible that the lower transformation from lycopene to carotenes contributed to the relatively stable content of lycopene.     

Development and validation of gene-based markers for shrunken2-Reference allele and their utilization in marker-assisted sweet corn (Zea mays Sachharata) breeding programme

The recessive shrunken2-Reference (sh2-Ref) allele is traditionally used in sweet corn cultivar development worldwide. However, non-availability of suitable gene-based marker(s) for sh2-Ref has considerably delayed its utilization in molecular breeding. Here, 7,320 bp sequence of Sh2 gene among five wild- and six sh2-Ref-based inbreds was analysed by employing 13 overlapping primers. SNP583 (A in wild and G in mutant) and SNP755 (T in wild and C in mutant) in 5’UTR, and SNP5112 (C in wild and T in mutant) in intron-12 clearly differentiated dominant (Sh2) and recessive (sh2-Ref) allele. These SNPs were used to develop four gene-based PCR markers. The newly developed markers were validated in six F₂ populations segregating for sh2-Ref allele, and 230 diverse inbreds of normal and sweet corn types. These markers were further used in genotyping of BC₁F₁ populations leading to successful selection of sh2-Ref allele in the marker-assisted breeding. Globally, this assumes great significance in molecular breeding of sweet corn.     

Effect of radiation intensity on the outcome of postharvest UV-C treatments

Studies on the use of UV-C radiation of fresh produce have focused on the selection of appropriate doses (energy per unit area) for different commodities, but little attention has been placed on the effect of radiation intensity (dose per unit time). In this study, tomatoes (Solanum lycopersicum cv. Elpida) and strawberries (Fragaria × ananassa cv. Camarosa), were harvested (breaker and 100% of surface red color respectively) and treated with 4 kJ m⁻² of UV-C, at low (3 W m⁻²) or high (33 W m⁻²) radiation intensities. Untreated fruits were used as controls. After the treatments and at different storage times the incidence of postharvest rots and the changes in fruit physical and chemical properties were determined. UV-C treatments reduced decay, with the effects being were more marked in fruit exposed to high intensities. Mold counts were unaffected by the treatments, suggesting that improved disease control did not result from greater germicide effect. In both fruit species exposure to UV-C radiation delayed ripening, evidenced as lower color development, pigment accumulation and softening. UV-C-treated fruit maintained better quality than the control. In strawberry, high intensity treatments were more effective to prevent deterioration than in tomato where the differences between UV-C treatments were subtler. Soluble solids, titratable acidity and ethanol soluble antioxidants were not affected regardless of the UV-C intensity. Consumer tests showed higher preference of fruit treated at high UV-C intensity. Results show that in addition to the applied dose, radiation intensity is a main factor determining the effectiveness of UV-C treatments and should not be over-sighted. For a given dose, increasing radiation intensity may in some cases maximize the benefits of UV-C on fruit quality, while significantly reducing the treatments time.     

Genetic mapping of quantitative trait loci (QTL) for resistance to septoria tritici blotch in a winter wheat cultivar Liwilla

Septoria tritici blotch (STB), caused by Mycosphaerella graminicola (anamorph Septoria tritici, syn. Zymoseptoria tritici), is present in most wheat-growing areas worldwide. Resistance breeding appears to be the most sensible approach to disease control. An attempt was made to identify loci associated with resistance to STB in a resistant winter wheat cultivar Liwilla. In the study we used a set of 74 doubled-haploid lines generated from anthers of F₁ hybrids between the resistant cultivar Liwilla and susceptible cultivar Begra. Four monopycnidiospore isolates of M. graminicola with diverse pathogenicity were used in tests on seedlings under controlled growth conditions and on adult plants under polytunnel conditions over a six year period. In both environments, the percentage leaf area covered by necrosis and covered by pycnidia were measured; time to heading and plant height were also recorded for the polytunnel experiments. Seven isolate-specific quantitative trait loci (QTLs) were associated with STB resistance: QStb.ihar-3A.2, QStb.ihar-6A, QStb.ihar-7A.2, QStb.ihar-1B, QStb.ihar-2B.2, QStb.ihar-3B, and QStb.ihar-5D. QTL on chromosome 5D and 7A represent novel STB resistance loci. The phenotypic variance explained by individual QTLs ranged from 9.5 % to 50.3 %. Three QTLs detected on chromosomes 3A, 7A and 1B showed major effects and were detected consistently in different environments. The locations of QStb.ihar-3A.2 and QStb.ihar-1B coincide with the resistance genes Stb6 and Stb11, respectively. Locus QStb.ihar-3B and a QTL for time to heading mapped to the same location, but are most likely not associated. Most of the mapped QTLs explain the resistance associated with both low necrosis and low pycnidia coverage.     

Resistance to Cephus cinctus Norton,the wheat stem sawfly,in a recombinant inbred line population of wheat derived from two resistance sources

Several species of wheat stem sawflies (WSS) are pests of wheat, including Cephus cinctus Norton in North America. Larvae feed inside the stems and cut the stem near plant maturity. The primary means of control is resistance due to solid stems, largely controlled by a locus on chromosome 3B (Qss.msub‐3BL). Cultivars that differed for WSS resistance, but with similar stem solidness, were crossed to determine the genetic basis for the differences. The cultivar ‘Scholar’ is susceptible, while ‘Conan’ shows resistance. Scholar and Conan possessed different alleles at Qss.msub‐3BL. Both alleles conferred solidness, yet the Conan allele conferred higher WSS resistance. An allele from Conan on chromosome 4A also decreased infestation and stem cutting. The 3B and 4A alleles from Conan acted in an additive fashion to provide increased WSS resistance without increasing stem solidness. Stem solidness has long been used by breeders as a proxy for WSS resistance because of its simplicity. Our results suggest that other resistance mechanisms may complement solid stems.     

Quantitative trait loci associated with lodging tolerance in soybean cultivar ‘Toyoharuka’

Lodging tolerance (LT) is an important trait for high yield and combine-harvesting efficiency in soybean [Glycine max (L.) Merr.]. Many previous studies have investigated quantitative trait loci (QTLs) for lodging score (LS) in soybean. Most of the investigated QTLs were located in the proximal region of maturity or growth habit loci. The aim of this study was to identify genetic factors for LT not associated with maturity or growth habit. QTL analysis was performed using a recombinant inbred line (RIL) population derived from a cross between ‘Toyoharuka’ (TH), a lodging-tolerant cultivar, and ‘Toyomusume’ (TM). The genotypes of TH and TM were estimated as both e1e2E3E4 and dt1. The average LS over 4 years was used for QTL analysis, identifying a major and stable QTL, qLS19-1, on chromosome 19. The LS of the near-isogenic line (NIL) with the TH allele at Sat_099, the nearest marker to qLS19-1, was significantly lower than the NIL with the TM allele at that position. The TH allele at Sat_099 rarely had a negative influence on seed yield or other agronomic traits in both NILs and the TM-backcrossed lines. Our results suggest that marker-assisted selection for qLS19-1 is effective for improving LT in breeding programs.     

Effect of 1-methylcyclopropene on ripening of ‘Canino’ apricots and ‘Royal Zee’ plums

‘Canino’ apricots and ‘Royal Zee’ plums were treated with 1000 nl l⁻¹ 1-methylcyclopropene (1-MCP) at 20 °C for 20 h following harvest before 0 °C storage. After 5 days storage for apricots and 10 days for plums and after 30 days storage for both, fruit were moved to 20 °C for ripening. In addition, apricots were stored for 20 days and then treated with 1-MCP concentrations of 10, 100 and 1000 nl l⁻¹ at removal and held for ripening. Ethylene production and respiration rate, as well as fruit quality of apricots varied with treatment. Ethylene production was efficiently inhibited by 1000 nl l⁻¹ 1-MCP in fruit treated after storage but not in fruit treated before storage. Fruit softening was associated with ethylene production and affected by 1-MCP in a concentration dependent manner when treated after storage, while 1-MCP did not affect softening in prestorage treated fruit. The color change of fruit was ethylene-independent and not affected by 1-MCP. Internal flesh browning was decreased by 1-MCP regardless of the concentration when treated after storage, while it was enhanced in fruit treated before storage. Decay development in apricots was decreased by 1-MCP in a concentration dependent manner. Ethylene production and respiration in ‘Royal Zee’ plums was greatly inhibited by 1-MCP during ripening after both short-term (10 day) and long term (30 day) storage. Parameters associated with ripening processes were decreased significantly by 1-MCP, including softening, color change, and loss of titratable acidity. These data demonstrate that 1-MCP has potential to delay ripening of apricots and plums, but the cultivar, maturity of fruit, and time of application must be chosen carefully. It is suggested that 1-MCP is more efficient for extending the shelf life and improving the quality of ‘Canino’ apricots directly marketed or after storage, whereas it might be a potent compound for extending both storage period and shelf life of ‘Royal Zee’ plums.     

Postharvest pigmentation in red Chinese sand pears (Pyrus pyrifolia Nakai) in response to optimum light and temperature

The development of red color in the peel of red Chinese sand pears (Pyrus pyrifolia Nakai) is influenced by temperature and light; however, the response patterns vary among different cultivars. In this study, we systematically investigated the influence of postharvest treatment with various temperatures (low, high, variant and constant) on detached mature fruit of red Chinese sand pear ‘Mantianhong’ and ‘Meirensu’. Fruit of red apple (Malus domestica Borkh.) ‘Royal Gala’ and red European pear (P. communis L.) ‘Cascade’ received the same treatments for comparison. Furthermore, the effects of light quality and irradiance level on ‘Mantianhong’ pears were evaluated at the optimum temperature for anthocyanin accumulation. Fruit firmness and concentrations of total soluble sugars and organic acids were measured to determine fruit quality. The effect of temperature on red Chinese sand pear fruit color was similar to that of apples, but not European pear. Moreover, low temperature more effectively induced red coloration in ‘Mantianhong’ and ‘Meirensu’ pears than high temperature; anthocyanin levels increased with increasing irradiance level from 0 to 532 μmol m⁻² s⁻¹, and UV-B and visible light synergistically improved the red color of the fruit. Therefore, a combination of low temperature and high intensity of UV-B/visible light could improve the postharvest coloration of red sand pear fruit. The results will contribute to an improved understanding of the mechanism responsible for the coloration of red Chinese sand pears and will aid development of new techniques to improve color in postharvest fruit.     

Inheritance of fruit flesh and skin colours in powdery mildew resistant cucumbers (Cucumis sativus L.)

Summary The flesh of ripe cucumber fruits can show different colours, varying from intense or dingy white to yellow and orange. These colours appear to be governed by two genes, for which the symbols V and W are proposed. If both genes are recessive, an orange colour is produced; if both are dominant they give a dirty white colour. No linkage with the genes for mildew resistance could be demonstrated.A very strong linkage was observed between mildew resistance and a dull green exterior fruit colour governed by one gene for which the symbol D is proposed.     

Characterization of ethylene biosynthesis and its regulation during fruit ripening in kiwifruit,Actinidia chinensis ‘Sanuki Gold’

Ethylene biosynthesis in kiwifruit, Actinidia chinensis ‘Sanuki Gold’ was characterized using propylene, an ethylene analog, and 1-methylcyclopropene (1-MCP), an inhibitor of ethylene perception. In fruit harvested between a young stage (66 days after pollination) (DAP) and an early commercial harvesting stage (143 DAP), 2 days of exposure to propylene were sufficient to initiate ethylene biosynthesis while in fruit harvested at commercial harvesting stage (154 DAP), 4 days of propylene treatment were required. This observation suggests that response of ethylene biosynthesis to propylene treatment in kiwifruit declined with fruit maturity. Propylene treatment resulted in up-regulated expression of AC-ACO1, AC-ACO2, AC-SAM1 and AC-SAM2, prior to the induction of AC-ACS1 and ethylene production, confirming that AC-ACS1 is the rate limiting step in ethylene biosynthesis in kiwifruit. Treatment of fruit with more than 5 μL L^?1 of 1-MCP after the induction of ethylene production subsequently suppressed ethylene production and expression of ethylene biosynthesis genes. Treatment of fruit with 1-MCP at harvest followed with propylene treatment delayed the induction of ethylene production and AC-ACS1 expression for 5 days. These observations suggest that in ripening kiwifruit, ethylene biosynthesis is regulated by positive feedback mechanism and that 1-MCP treatment at harvest effectively delays ethylene production by 5 days.     

Identification of the fourth allele of the ANS (anthocyanidin synthase) gene and its effect on red color intensity in onions (Allium cepa)

Bulb color in onions (Allium cepa) is an important trait, and homogenous red coloration is desirable in red onion cultivars. The gene encoding anthocyanin synthase (ANS) is required for anthocyanin biosynthesis in onions. We have previously described three different alleles of the ANS gene. Here we report identification of the fourth allele of ANS, ANS-h1, found in a dark red doubled haploid line. ANS-h1 is similar to a non-functional allele found in Brazilian yellow cultivars except that it has several point mutations and indels throughout the promoter and coding regions, none of which are predicted to inactivate enzymatic activity. F₂ and backcross populations originating from the crosses between wild-type (ANS-L) allele-containing red and pink (ANS-p) allele-containing white or yellow parents show a discrete segregation ratio of 3 red to 1 light pink, indicating that the wild-type allele is completely dominant over the pink allele. In contrast, segregating populations derived from the crosses between ANS-h1 allele-containing red and the same white or yellow parents show a gradient of red intensity from light pink to dark red, suggesting that other genetic factors may affect expression of ANS-h1. A newly developed PCR-based marker and two previously developed markers for allelic selection of the ANS gene were used to examine allele composition in fifty-six breeding lines and commercial cultivars. Most lines are heterogeneous for the ANS gene with two or three alleles detected. The frequency of the pink allele is low in red breeding lines, but it is predominant in white and yellow lines.     

Ethylene biosynthesis in apricot: Identification of a ripening-related 1-aminocyclopropane-1-carboxylic acid synthase (ACS) gene

Apricots are climacteric fruits with a high susceptibility to flesh softening and loss of flavor during postharvest storage, and most of the ripening processes are regulated by ethylene, which also has an effect on its own biosynthesis. To understand this process in apricot, inhibition of ethylene biosynthesis and perception was performed for studying key genes involved in the ethylene biosynthetic pathway. Apricots, cv. “Patterson”, were harvested with yellow-green ground color and immediately treated with either the ethylene perception inhibitor 1-methyl cyclopropene (1-MCP) at 10 μL L⁻¹ or the ethylene biosynthesis inhibitor aminoethoxyvinylglycine (AVG) at 1 g L⁻¹. After treatment, quality and physiological attributes such as firmness, color, total soluble solids, acidity, fruit weight, ethylene production and respiration rates were evaluated every 2 d until they ripened at 20 °C. Gene expression analysis was performed by quantitative polymerase chain reaction (qPCR). Both ethylene inhibitors were effective in reducing ethylene production, respiration rate and fruit softening. Three 1-aminocyclopropane-1-carboxylic-acid synthase (ACS) genes were characterized, but only the expression of ACS2 was highly reduced by ethylene inhibition, suggesting a key role in ethylene synthesis at ripening. Contrarily, ACS1 and ACS3 showed a higher expression under ethylene inhibition suggesting that the corresponding genes are individually regulated in a specific mode as observed in other climacteric fruits. Finally, changes in 1-aminocyclopropane-1-carboxylic-acid oxidase genes did not show a consistent pattern of ethylene modulation.     

A pubescence color gene enhances tolerance to cold-induced seed cracking in yellow soybean

In yellow soybean, severe cold weather causes seed cracking on the dorsal side. Yellow soybeans carry the I or iⁱ allele of the I locus and have a yellow (I) or pigmented (iⁱ) hilum. We previously isolated an additional allele, designated as Ic, of the I locus, and reported that yellow soybeans with the IcIc genotype may be tolerant to cold-induced seed cracking. The Ic allele by itself, however, does not confer high tolerance. The association of a pubescence color gene (T) with suppression of low-temperature-induced seed coat deterioration has been previously reported. In the present study, we tested whether T is effective for the suppression of cold-induced seed cracking using two pairs of near-isogenic lines for the T locus in the iⁱiⁱ or IcIc background. In both backgrounds, the cracked seed rate of the near-isogenic line with the TT genotype was significantly lower than that with the tt genotype, which indicates that T has an inhibitory effect on cold-induced seed cracking. Furthermore, we also showed that gene pyramiding of Ic and T can improve tolerance to cold-induced seed cracking. Our findings should aid the development of highly SC-tolerant cultivars in soybean breeding programs.