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Cucurbit Genetics Cooperative

Gene List for Squash


Harry S. Paris
A.R.O., Newe Ya'ar Research Center, Ramat Yishay 30-095 (Israel)

Eileen Kabelka
University of Florida, Gainesville, Fl 32611 (U.S.A.)

The genus Cucurbita L. contains 12 or 13 species (50).  As far as is known, all have a complement of 20 pairs of chromosomes (2n = 40).

This gene list for Cucurbita contains detailed sources of information, being modeled after the one for cucumber presented by Wehner and Staub (103) and its update by Xie and Wehner (109). In order to more easily allow confirmation of previous work and as a basis for further work, information has been included concerning the genetic background of the parents that had been used for crossing.  Thus, in addition to the species involved, the cultivar-group (for C. pepo), market type (for C. maxima, C. moschata), and/or cultivar name are included in the description wherever possible.

Genes affecting phenotypic/morphological traits are listed in Table 1. The data upon which are based identifications and concomitant assignment of gene symbols vary considerably in their content.  No attempt is made here to assess the certainty of identifications, but gene symbols have been accepted or assigned only for cases in which at least some data are presented. The genes that are protein/isozyme variants are listed in Table 2. It can be seen from Tables 1 and 2 that approximately 70 genes have been identified for C. pepo L, for C. moschata Duchesne 25 and for C. maxima Duchesne 19. For the interspecific cross of C. maxima × C. ecuadorensis Cutler & Whitaker, 29 genes have been identified, of which 25 are isozyme variants.  A few genes have also been identified in four of the wild species (C. okeechobeensis Bailey, C. lundelliana Bailey, C. foetidissima HBK and C. ecuadorensis) and in several other interspecific crosses.

Some genes are listed as occurring in more than one species.  This does not necessarily indicate that these genes reside at identical locations in the genome of different species.

New additions to the list of Cucurbita genes include a number of omissions as well as a number of new genes published after the last update.  Those that had been omitted are three unnamed genes for fruit bitterness (3). They are herein designated Bitter fruit-1, -2, and -3, symbols Bi-1, Bi-2, and Bi-3. This has necessitated the modification of the symbols for the two previously identified genes (12, 30, 32) for Bitter fruit as Bimax and Bi-0.  Newly identified genes that have been published since the last update are: ae (androecy enhancer), Crr-1, Crr-2, and Crr-3 (Crown rot resistance-1, -2, and -3), gl-2 (glabrous-2), and l-2R (light type-2 Reverse striping). The symbols ae, Crr-1, Crr-2, and Crr-3 are herein assigned for the first time. Before choosing a gene name and symbol, researchers are urged to consult this Gene List as well as the rules of Gene Nomenclature for the Cucurbitaceae that appears near the end of this Cucurbit Genetics Cooperative Report in order to avoid confusion arising from duplication of gene names and symbols. Please contact us if you find omissions or errors in this Gene List.

Several cases of genetic linkage have been reported: D – mo-2 (61) and M – Wt (C. pepo) (72) and Bi – Lo-2 (C. ecuadorensis × C. maxima) (32). Some of the isozyme variants observed by Weeden & Robinson (102) were also found to be linked to one another. RAPD markers have been categorized and organized into linkage groups and are not listed here but can be found in Brown and Myers (5) and Zraidi and Lelley (111). These two maps cannot be easily compared, as they were constructed using different mapping populations; RAPD markers are population-specific. Neither map gives complete coverage of the Cucurbita genome. Both maps contain morphological traits, either as single genes or as quantitative trait loci (QTLs), which are listed in Table 3. More recently, a map for Cucurbita pepo has been constructed using RAPDs, AFLPs, and SSRs (27, 113). Over 300 markers were mapped, with coverage of some 2,200 cM of the genome, 20 linkage groups and a map density of 2.9 cM.

Sequenced genes can be valuable to breeders and geneticists, as the differences in the gene sequences that result in the phenotypes of interest can be used in marker-assisted selection. Unlike random markers, these gene-specific, allele-specific markers are completely linked to the genes of interest. Most of the genes sequenced in Cucurbita have been isolated by researchers doing comparative studies of specific genes across plant families; usually only a single allele is available. Nonetheless, we have included a list of the sequenced genes as Table 4 because the sequences could be useful as a starting point for breeders interested in isolating the genes from lines of differing phenotype. In addition to the genes listed here, there exists a collection of partial sequences from mRNA for genes differentially expressed during seed development in C. pepo. These expressed sequence tags were identified in a study of the naked seed trait. The Gene Accession numbers for these sequences are CD726806 through CD726832.

Table 1.  Phenotypic/Morphological Characteristics

Gene Symbol




Preferred Synonym Character Species Reference(s)
a   androecious.  Found in ‘Greckie’; produces only male flowers, recessive to A. pepo 41
ae*   androecy enhancer. From cross between two vegetable-marrow cultivars, the strongly male ‘Vegetable Spaghetti’,  ae/ae , and ‘Bolognese’, Ae/Ae. pepo 48
B   Bicolor. Precocious yellow fruit pigmentation; pleiotropic, affecting fruit and foliage, modified by Ep-1, Ep-2 and Ses-B.  Originally from ‘Vaughn’s Pear Shaped’ ornamental gourd. B in C. moschata ‘Precocious PI 165561’ derived from C. pepo through backcrossing. Complementary to L-2 for intense orange, instead of light yellow, fruit-flesh color. pepo, moschata 57, 74, 84, 91, 93
Bmax B-2 Bicolor. Precocious yellow fruit pigmentation, from subsp. andreana PI 165558 maxima 92, 95
Bimax* Bi Bitter fruit. High cucurbitacin content in fruit.  Bi from C. maxima subsp. andreana and C. ecuadorensis; bi from C. maxima subsp. maxima, including ‘Queensland Blue’.  Linked to Lo-2 maxima, maxima × ecuadorensis 12, 32
Bi-0 * Bi Bi-0 from wild Texan gourd; bi-0 from zucchini squash. Might be identical with either Bi-1 or Bi-2. pepo 30
Bi-1 *   In cross of C. pepo ×C. argyrosperma, three complementary dominant alleles are needed for bitterness. Bi-1 from C. pepo straightneck ‘Goldbar’, bi-1 from C. argyrosperma ‘Green Striped Cushaw’. pepo × argyrosperma 3
Bi-2 *   In cross of C. pepo ×C. argyrosperma, three complementary dominant alleles are needed for bitterness. Bi-2 from C. pepo straightneck ‘Goldbar’, bi-2 from C. argyrosperma ‘Green Striped Cushaw’. pepo × argyrosperma 3
Bi-3 *   In cross of C. pepo ×C. argyrosperma, three complementary dominant alleles are needed for bitterness. Bi-3 from C. argyrosperma ‘Green Striped Cushaw’, bi-3 from C. pepo straightneck ‘Goldbar’. pepo × argyrosperma 3
bl   blue fruit color. Incompletely recessive to Bl forgreen fruit color, in hubbard squash. maxima 33
Bn   Butternut fruit shape, from ‘New Hampshire Butternut’, dominant to bn for crookneck fruit shape, as in ‘Canada Crookneck’. moschata 52
Bu D Bush habit.  Short internodes; dominant to vine habit, bu, in young plant stage.  In C. pepo, Bu in ‘Giant Yellow Straightneck’ and near-isogenic line of ‘Table Queen’, bu in ‘Table Queen’ acorn.  In C. maxima, Bu from inbred line, bu from ‘Delicious’. In C. moschata, Bu from inbred line, bu from undisclosed parent. pepo, maxima, moschata 18, 31, 90, 106
Cmv   Cucumber mosaic virus resistance, from Nigerian Local.  Dominant to cmv for susceptiblity, from ‘Waltham Butternut’. moschata 4
cr   cream corolla.  Cream to nearly white petals, cr from C. okeechobeensis; Cr from C. moschata ‘Butternut’ incompletely dominant (yellow petals for Cr/cr, and orange for Cr/Cr) moschata × okeechobeensis 81
Crr-1*   Crown rot resistance. Resistance to Phytophthora capsici, introgressed from C. lundelliana and C. okeechobeensis subsp. okeechobeensis into a breeding line of C. moschata. One of three complementary dominant genes for resistance. Genotype of the susceptible C. moschata ‘Butterbush’ is crr-1/crr-1. moschata 56
Crr-2*   Crown rot resistance. Resistance to Phytophthora capsici, introgressed from C. lundelliana and C. okeechobeensis subsp. okeechobeensis into a breeding line of C. moschata, One of three complementary dominant genes for resistance. Genotype of the susceptible C. moschata ‘Butterbush’ is crr-2/crr-2. moschata 56
Crr-3*   Crown rot resistance. Resistance to Phytophthora capsici, introgressed from C. lundelliana and C. okeechobeensis subsp. okeechobeensis into a breeding line of C. moschata. One of three complementary dominant genes for resistance. Genotype of the susceptible C. moschata ‘Butterbush’ is crr-3/crr-3. moschata 56
cu   cucurbitacin-B reduced; cu for reduced cucurbitacin-B content of cotyledons of ‘Early Golden Bush Scallop’; Cu for high cucurbitacin content of cotyledons of ‘Black Zucchini’. pepo 89
D   Dark stem.  Series of three alleles observed in C. pepo: D for dark stem and dark intermediate-age fruit, Ds for dark stem but fruit not affected, and d for light stem and fruit not affected, with dominance D > Ds > dD from ‘Fordhook Zucchini’, Ds from ‘Early Prolific Straightneck’; d from ‘Vegetable Spaghetti’. Epistatic to genes l-1 and l-2 when either is homozygous recessive; linked to mo-2.  In C. maxima, only the fruit was observed: D for dark intermediate-age fruit from the zapallito ‘La Germinadora’; d for light intermediate-age fruit from a variant zapallito breeding stock. pepo, maxima 26, 45, 60, 61, 64, 73, 86
de   determinate plant habit; stem lacking tendrils and terminating with female flowers.  Recessive to De for indeterminate plant habit.  De from ‘Jeju’ and ‘Sokuk’, de from inbred designated “Det”. moschata 42
Di   Disc fruit shape.  From scallop squash, dominant to spherical or pyriform. pepo 97, 104
Ep-1   Extender of pigmentation-1; modifier of BEp-1 incompletely dominant to ep-1 and additive with Ep-2Ep-1 from ‘Small Sugar 7 × 7’ pumpkin; ep-1 from ‘Table King’ acorn. pepo 96
Ep-2   Extender of pigmentation-2; modifier of BEp-2 incompletely dominant to ep-2 and additive with Ep-1Ep-2 from ‘Table King’ acorn; ep-2 from ‘Small Sugar 7 × 7’ pumpkin. pepo 96
Fr   Fruit fly (Dacus cucurbitae) resistance. Fr from ‘Arka Suryamukhi’, dominant to fr forsusceptibility. maxima 53
fv   fused vein. Fusion of primary leaf veins, subvital male gametophyte; found in hull-less-seeded pumpkin breeding line. pepo 8, 9
G a, m Gynoecious sex expression; dominant to g for monoecious sex expression. foetidissima 19, 24
Gb   Green band on inner side of base of petal, from a scallop squash; dominant to gb, for no band, from a straightneck squash. pepo 20
gc   green corolla.  Green, leaf-like petals, sterile; in unspecified F2 population. pepo 99
gl-1*   gl glabrous, lacking trichomes maxima 37
gl-2   glabrous, lacking trichomes; gl-2 mutant found in straightneck squash pepo 108
Gr G Green rind. Dominant to buff skin of mature fruit.  Gr from ‘Long Neapolitan’, gr from ‘Butternut’. moschata 77
grl   gray leaf. Recessive to green leaf.  Recessive grl derived from cross of  zapallito-type line of C. maxima and a butternut-type line of C. moschata.  Dominant Grl from zapallito-type C. maxima. maxima × moschata 44
Hi   Hard rind inhibitorHi, for hard-rind inhibition, from C. maxima ‘Queensland Blue’; hi, for no hard-rind inhibition, from C. ecuadorensis. maxima × ecuadorensis 30
Hr   Hard rind. Hr for hard (lignified) rind inornamental gourd, straightneck squash, and zucchini; hr for soft (non-lignified) rind in ‘Small Sugar’ pumpkin and ‘Sweet Potato’ (‘Delicata’).  Complementary to Wt for Warty fruit. pepo 47, 85
i   intensifier of the cr gene for cream flowers.  Cr/__ I/__ for intense orange or yellow flowers, Cr/__ i/i for light orange or yellow flowers, cr/cr I/__ for cream flowers, cr/cr i/i for white flowers.  I from C. moschata ‘Butternut’, i from C. okeechobeensis. moschata × okeechobeensis 81
I-mc Imc Inhibitor of mature fruit color; dominant to i-mc for no inhibition. I-mc in a scallop squash. pepo 10
I-T   Inhibitor of the T gene for trifluralin resistance.  I-T from ‘La Primera’; i-t from ‘Ponca’ and ‘Waltham Butternut’. moschata 1
l-1 c, St light fruit coloration-1. Light intensity of fruit coloration. Series of five alleles observed in C. pepo which, in complementary interaction with the dominant L-2 allele, give the following results: L-1 for uniformly intense/dark fruit coloration, from ‘Fordhook Zucchini’; l-1BSt for broad, contiguous intense/dark stripes, from ‘Cocozelle’; l-1St for narrow, broken intense/dark stripes, from ‘Caserta’; l-1iSt for irregular intense/dark stripes, from ‘Beirut’ vegetable marrow; l-1 for light coloration, from ‘Vegetable Spaghetti’, with dominance of L-1 > (l-1BSt > l-1St) ≥ l-1iSt > l-1. In C. maxima, L-1 from the zapallito ‘La Germinadora’; l-1 from a variant zapallito breeding stock. pepo, maxima 26, 45, 62, 67, 63, 67, 73, 82, 91
l-2 r light fruit coloration-2. Light intensity of fruit coloration. Series of four alleles observed in C. pepo, which, in complementary interaction with dominant alleles at the l-1 locus, give the following results: L-2 for intense/dark fruit coloration, with L-1 from ‘Fordhook Zucchini’ and intense/dark fruit stripes, with l-1BSt from ‘Cocozelle’; allele L-2w has delayed and weaker effect than L-2, from C. pepo subsp. fraterna; l-2R confers reversal of color, that is, stripes lighter than the background in combination with any of the striping alleles at the l-1 locus, or completely light fruit in the presence of L-1, from C. pepo subsp. texana ‘Delicata’; l-2 for light coloration, from ‘Vegetable Spaghetti’, with dominance of L-2 > L-2w > l-2. Dominant L-2 is also complementary with B for intense orange, instead of light yellow, fruit-flesh color and with recessive qi for intense exterior color of young fruit. In C. maxima, L-2 from the zapallito ‘La Germinadora’; l-2 from a variant zapallito breeding stock.  pepo, maxima 26, 45, 57, 65, 68, 69, 73
lo-1 l lobed leaves-1; recessive to Lo-1 for non-lobed leaves maxima 21
Lo-2   Lobed leaves-2. Lo-2 for lobed leaves in C. ecuadorensis dominant to lo-2 for unlobed leaves in C. maxima. Linked to Bi. ecuadorensis × maxima 32
lt   leafy tendril. Tendrils with laminae; lt found in ornamental gourd. pepo 83
ly   light yellow corolla. Recessive to orange yellow; ly found in ornamental gourd. pepo 83
M   Mottled leaves.  M for silver-gray areas in axils of leaf veins, dominant to m for absence of silver-gray.   For C. maxima, M in ‘Zuni’ and m in ‘Buttercup’ and ‘Golden Hubbard’. For C. pepo, M in ‘Caserta’ and inbred of ‘Striato d’Italia’ cocozelle; m in ‘Early Prolific Straightneck’ and ‘Early Yellow Crookneck’. For C. moschata, M in ‘Hercules’ and ‘Golden Cushaw’, m in butternut type. Weakly linked to Wt. pepo, maxima, moschata 14, 66, 82, 87
Mldg   Mottled light and dark green immature fruit color; germplasm unspecified. Dominant to mldg for non-mottled. moschata 6
mo-1   mature orange-1; complementary recessive gene for loss of green fruit color prior to maturity.  Mo-1 from ‘Table Queen’ acorn; mo-1 from ‘Vegetable Spaghetti’. pepo 61
mo-2   mature orange-2; complementary recessive gene for loss of green fruit color prior to maturity.  Mo-2 from ‘Table Queen’ acorn; mo-2 from ‘Vegetable Spaghetti’.  Linked to D. pepo 61
ms-1 ms1 male sterile-1. Male flowers abort before anthesis, derived from a cross involving ‘Golden Hubbard’, recessive to Ms-1 for male fertile. maxima 88
ms-2 ms2 male sterile-2. Male flowers abort, sterility expressed as androecium shrivelling and turning brown; ms-2 from ‘Eskandarany’ (PI 228241). pepo 23
ms-3 ms-2 male sterile-3. maxima 39
m-zymmos   modifier of dominance of zucchini yellow mosaic virus resistance; confers resistance to otherwise susceptible Zymmos/zymmos heterozygotes. M-zymmos in ‘Soler’, m-zymmos in ‘Waltham Butternut’ and ‘Nigerian Local’. moschata 55
n h naked seeds.  Lacking a lignified seed coat, n from oil-seed pumpkin. pepo, moschata 29, 86, 107, 112, 113
pl   plain light fruit color, pl from ‘Beirut’ vegetable marrow and ‘Fordhook Zucchini’; Pl in ‘Vegetable Spaghetti’. pepo 58
Pm   Powdery mildew resistance.  Resistance to Podosphaera xanthii; Pm from C. lundelliana. lundelliana 13, 70
Pm-0   Powdery mildew resistance. Resistance to Podosphaera xanthii; Pm-0 from C. okeechobeensis and in C. pepo. okeechobeensis, pepo 11, 37
pm-1   powdery mildew resistance in C. moschata. Series of three alleles: pm-1P for susceptibility from ‘Ponca’ dominant to pm-1L for resistance from ‘La Primera’, which is dominant to pm-1W for susceptibility in ‘Waltham Butternut’. moschata 2
pm-2   powdery mildew resistance in C. moschata ‘Seminole’, recessive to Pm-2 for susceptibility moschata 2
prv   papaya ringspot virus resistance, in Nigerian Local, recessive to Prv for susceptibility, in ‘Waltham Butternut’. moschata 4
qi   quiescent intense.  Recessive to Qi for not intense and complementary to L-2 for intense young fruit color; little or no effect on mature fruit. Qi from ‘Vegetable Spaghetti’; qi from ‘Jack O’Lantern’ pumpkin and ‘Verte non-coureuse d’Italie’ cocozelle. pepo 63, 66
Rd   Red skin. Red external fruit color; dominant to green, white, yellow and gray. Rd from ‘Turk’s Cap’; rd from ‘Warted Hubbard’. maxima 46
ro   rosette  leaf.  Lower lobes of leaves slightly spiraled, ro derived from an ornamental gourd. pepo 47
s-1 s sterile. Male flowers small, without pollen; female flower sterile.  Derived from crossing ‘Greengold’ with ‘Banana’. maxima 34
s-2   sterile.  Male flowers small, without pollen and female flower sterile; mutant in powdery mildew resistant, straightneck squash breeding line. pepo 7
Ses-B   Selective suppression of gene B. Suppression in foliage of precocious yellowing conferred by B. Ses-B in straightneck breeding line dominant to ses-B in ‘Jersey Golden Acorn’. pepo 94
sl   silverleaf resistance.  Recessive to Sl for susceptibility. In C. moschata, Sl from ‘Soler’; sl from PI 162889 and butternut types. In C. pepo, Sl from ‘Black Beauty’ zucchini and sl from Zuc76 breeding line. moschata, pepo 28, 110
slc   Squash leaf curl virus resistance; derived from C. moschata. pepo 50
sp   spaghetti flesh, breaking into strands after cooking pepo 49
T   Trifluralin resistance.  Dominant to susceptibility to the herbicide; modified by I-TT in ‘La Primera’; t in ‘Ponca’ and ‘Waltham Butternut’. moschata 1
uml   umbrella-like; leaves shaped like partially opened umbrella. Recessive uml derived from a cross of C. maxima ‘Warzywna’ and a C. pepo inbred; dominant Uml from ‘Warzywna’.  maxima × pepo 75
v   virescent. Yellow-green young leaves, v found in ‘Golden Delicious’. maxima 22
W   Weak fruit coloration.  Dominant to w for intense-pigmented mature fruit; W from scallop squash. Complementary to Wf for white external fruit color. pepo 59, 91, 97
wc   white corolla.  Derived from ‘Ispanskaya’ × ‘Emerald’.  Recessive to Wc for normal orange-yellow corolla maxima 40
Wf   White flesh. Dominant to wf for colored flesh.  Wf in a scallop squash, wf in a straightneck squash. Complementary to W for white external fruit color. pepo 20, 59, 97
Wmv   Watermelon mosaic virus resistance.  From “Menina” and “Nigerian Local”, dominant to wmv for susceptibility in ‘Musquée de Provence’ and ‘Waltham Butternut’. May be linked with or identical to Zym-1. moschata 4, 24
Wmvecu   Watermelon mosaic virus resistance. From C. ecuadorensis, in a cross with an unspecified C. maxima. maxima x ecuadorensis 95
Wt   Warty fruit. Dominant to non-warted, wt, and complementary to Hr, with fruit wartiness being expressed only in the presence of the dominant Hr allele. Wt in straightneck, crookneck, and ‘Delicata’; wt in zucchini, cocozelle, and ‘Small Sugar’ pumpkin.  Weakly linked to M. pepo 66, 79, 97
wyc   white-yellow corolla; isolated in ‘Riesen-Melonen’.  Recessive to Wyc for normal orange-yellow corolla. maxima 40
Y   Yellow fruit color. Y for yellow fruit color of intermediate-age fruits, from straightneck and crookneck squash, dominant to y for green intermediate-age fruit color, from vegetable marrow, ornamental gourd, and cocozelle. pepo 72, 82, 90, 91, 97
yg   yellow-green leaves and stems maxima 39
Ygp   Yellow-green placenta. Dominant to yellow placental color.  Ygp in a scallop squash, ygp in a straightneck squash. pepo 20
ys   yellow seedling. Lacking chlorophyll; lethal pepo 47
zymecu   zucchini yellow mosaic virus resistance, recessive to susceptibility; zymecu from C. ecuadorensis, Zymecu from C. maxima ‘Buttercup’. ecuadorensis 80
zymmos   zucchini yellow mosaic virus resistance, recessive to susceptibility; zym?mos from ‘Soler’, Zymmos from ‘Waltham Butternut’. moschata 55
Zym-0   Zucchini yellow mosaic virus resistance.  Zym-0 from C. moschata ‘Nigerian Local’ dominant to zym-0 for susceptibility from ‘Waltham Butternut’. Perhaps one of two separate genes for resistance in ‘Nigerian Local’. moschata 4, 47, 55
Zym-1   Zucchini yellow mosaic virus resistance. Zym-1 from C. moschata ‘Menina’dominant to zym-1 forsusceptibility from C. moschata ‘Waltham Butternut’.  Zym-1 transferred via backcrossing to C. pepo ‘True French’ zucchini, in which it confers resistance through complementary interaction with Zym-2 and Zym-3. Zym-1 is either linked with Wmv or also confers resistance to watermelon mosaic virus. moschata, pepo 24, 55, 70, 71 
Zym-2   Zucchini yellow mosaic virus resistance-2.  Dominant to susceptibility and complementary to Zym-1Zym-2 from C. moschata ‘Menina’. Zym-2 in C. pepo derived from C. moschata, in near-isogenic resistant line of ‘True French’ zucchini; zym-2 from C. pepo ‘True French’. moschata, pepo 70
Zym-3   Zucchini yellow mosaic virus resistance-3.  Dominant to susceptibility and complementary to Zym-1.  Zym-3 from C. moschata ‘Menina’. Zym-3 in C. pepo derived from C. moschata, in near-isogenic resistant line of ‘True French’ zucchini; zym-3 from C. pepo ‘True French’. moschata, pepo 70

*Proposed new gene symbol.


Table 2. Isozyme Variants

Gene Symbol





Preferred Synonym No. alleles observed Character Species Reference(s)
Aat-1 Aat 8 Aspartate aminotransferase-1. Variant among accessions. pepo 17, 36
Aat-3   2 Aspartate aminotransferase-3.  Variant among wild populations. pepo 17
Aat-4   3 Aspartate aminotransferase-4.  Variant  among wild populations. pepo 17
Aat-mb   2 Aspartate aminotransferase – microbody maxima x ecuadorensis 102
Aat-m1   2 Aspartate aminotransferase mitochondria-1 maxima x ecuadorensis 102
Aat-m2   2 Aspartate aminotransferase mitochondria-2 maxima x ecuadorensis 102
Aat-p2   2 Aspartate aminotransferase plastid-2 maxima x ecuadorensis 102
Acp-1   2 Acid phosphatase-1 maxima x ecuadorensis 102
Acp-2   2 Acid phosphatase-2 maxima x ecuadorensis 102
Adh   2 Alcohol dehydrogenase pepo 105
Aldo-p   2 Aldolase – plastid maxima x ecuadorensis 101
Est-1 Est 2 Esterase maxima x ecuadorensis 100, 102
Gal-1   2 b-galactosidase-1 maxima x ecuadorensis 102
Gal-2   2 b-galactosidase-2 maxima x ecuadorensis 102
G2d-1   3 Glycerate dehydrogenase-1. Variant among wild populations. pepo 17
G2d-2   2 Glycerate dehydrogenase-2. Variant among wild populations. pepo 17
Got-1   5 Glutamine oxaloacetate-1. Variant among accessions, wild populations, and among Cucurbita species. pepo 15, 16, 38, 105
Got-2   3 Glutamine oxaloacetate-2. Variant among species. maxima x ecuadorensis 102
Gpi   2 Glucosephosphate isomerase.  Variant among accessions. pepo 36
Gpi-3   2 Glucosephosphate isomerase-3.  Variant among wild populations. pepo 17
Gpi-c1   2 Glucosephosphate isomerase cytosolic-1 maxima x ecuadorensis 102
Gpi-c2   2 Glucosephosphate isomerase cytosolic-2 maxima x ecuadorensis 102
Idh-1   4 Isocitrate dehydrogenase-1.  Variant among accessions, wild populations, and Cucurbita species. pepo 15, 16, 17, 38, 105
Idh-2   2 Isocitrate dehydrogenase-2. Variant among accessions, wild populations, and Cucurbita species.  pepo 15, 16, 17, 38, 105
Idh-3   2 Isocitrate dehydrogenase-3. Variant among accessions and populations. pepo 15, 16, 17, 38
Lap-1 Lap 4 Leucine aminopeptidase. Variant among C. pepo accessions. maxima x ecuadorensis; pepo 17, 36, 100, 102
Mdh-1 Mdh 7 Malate dehydrogenase. Variant among accessions. pepo 36
Mdh-2   3 Malate dehydrogenase-2. Variant among accessions, wild populations, and Cucurbita species. pepo 15, 16, 17, 38, 105
Mdh-3   3 Malate dehydrogenase-3. Variant among accessions, wild populations, and Cucurbita species. pepo 15, 16, 17, 38, 105
Mdh-m1   2 Malate dehydrogenase mitochondria-1 maxima x ecuadorensis 102
Mdh-m2   2 Malate dehydrogenase mitochondria-2 maxima x ecuadorensis 102
Mdh-c2   2 Malate dehydrogenase cytosolic-2 maxima x ecuadorensis 102
Per-1   2 Peroxidase-1 maxima x ecuadorensis 102
Per-2   3 Peroxidase-2.  Variant among accessions and wild populations. pepo 15, 16, 38
Per-3   2 Peroxidase-3 maxima x ecuadorensis 102
Pgi-1   2 Phosphoglucase isomerase-1 pepo 15
Pgi-2   2 Phosphoglucase isomerase-2. Variant among Cucurbita species. pepo 15, 38, 105
Pgi-3   4 Phosphoglucase isomerase-3. Variant among accessions, wild populations, and Cucurbita species. pepo 15, 16, 38, 105
Pgm-1 Pgm 2 Phosphoglucomutase.  Variant among accessions. pepo 36
Pgm-2   4 Phosphoglucomutase-2. Variant among accessions, wild populations, and Cucurbita species. pepo 15, 16, 38, 105
Pgm-5   2 Phosphoglucomutase-5. Variant among wild populations. pepo 17
Pgm-6   2 Phosphoglucomutase-6. Variant among wild populations. pepo 17
Pgm-c2   2 Phosphoglucomutase cytosolic-2 maxima x ecuadorensis 102
Pgm-p   2 Phosphoglucomutase plastid maxima x ecuadorensis 102
Skd-1   6 Shikimate dehydrogenase. Variant among wild populations. pepo 17
Skdh   5 Shikimate dehydrogenase. Variant among C. pepo accessions. maxima x ecuadorensis; pepo 36, 102
Sod-1   2 Superoxide dismutase-1 maxima x ecuadorensis 102
Tpi-c2   2 Triosephosphatase isomerase cytosolic-2 maxima x ecuadorensis 102
Tpi-p2   2 Triosephosphatase isomerase plastid-2 maxima x ecuadorensis 102


Table 3. Mapped Phenoypic/Morphological Characteristics

Trait Symbol Linked Marker(s) Recombination Distance (cM) Reference(s)
Precocious yellow fruit B I10_1700 27.1 5
Bush growth habit Bu CMTp131 7.8 27
Dwarf Bu S1225_548, SCAR3_398 2.29 43
Leaf Mottle M H14_600
Seed Coat n AK11_340 4.4 111
Hull-less seed n CMTp58, CMTp151, CMTm115, CMTm239 1.5 - 3.6 27
Mature Fruit Color [none given] G17_700 9.7 5
Fruit Length (QTL) AE07_165, AC10_490, AJ20_420, P13_750, J01_600, AO20_1200, T08_460, AB08_540, AE09_1600   111
Fruit Width (QTL) AE07_165, AJ20_420, AM10_950, AG08_440   111
Fruit Length/width Ratio (QTL) AE07_165, AC10_490, AJ20_420, P13_750, J01_600   111
No. of Fruit Chambers (QTL) P13_950, AE08_470   111
Leaf Indentation (QTL) F10_400, K11_950, G2_400   5
Fruit Shape (QTL) F8_1050, B8_900, H19_500   5


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Table 4. Genes with known DNA sequence.

Gene Symbol* Gene Accession (Putative) Function Source Ref.
AIG-2 AY666083 aspartic protease inhibitor C. maxima **
PRB1 AY326308 phloem RNA-binding protein C. maxima ‘Big Max’ **
GAIP AY32630, AY326307 gibberellic acid insensitive phloem protein (two very similar genes) C. maxima ‘Big Max’ **
FAD2 AY525163 omega-6 fatty acid desaturase C. pepo zucchini **
NIP1 AJ544830 Nod26-like protein C. pepo zucchini 35
PP2 AY312402 phloem protein 2 lectin (includes promoter region) C. moschata crookneck **
PP2 AF150627 phloem protein 2 lectin C. moschata crookneck **
PP2 Z22647 phloem protein 2 lectin C. pepo ‘Autumn Gold’ 61
PP2 Z17331 phloem protein 2 lectin C. maxima ‘Big Max’ 5
PP2 L31550, L31551, L31552 phloem protein 2 (three alleles) C. maxima **
GA2OX, GA20OX, GA3OX AJ315663, AJ302041, AJ308480, AJ302040 gibberellin oxidases (two sequences for GA2OX) C. maxima ‘Riesenmelone’ **
  U61385 gibberellin 20-oxidase C. maxima ‘Riesenmelone’ 38
  U63650 gibberellin 2 beta,3 beta hydroxylase C. maxima ‘Riesenmelone’ 39
  AJ006453 gibberellin 3 beta hydroxylase C. maxima ‘Riesenmelone’ **
  U61386 gibberellin dioxygenase C. maxima ‘Riesenmelone’ 37
Moschatin 1 through 5 AF462349, AF504011, AY25646, AY27921, AY279217 ribosome-inactivating protein C. moschata crookneck **
CPS1 AB109763 copalyl diphosphate synthase; gibberellin biosynthesis C. maxima **
CPS AF049905, AF049906 copalyl diphosphate synthase; gibberellin biosynthesis (2 genes) C. maxima 55
Hsc70 AF527794, AF527795, AF527796 cell-autonomous heat shock protein; chaperonin 70 (multiple sequences) C. maxima 1
  AB061204 thioredoxin h C. maxima **
Puga, Pugb, Pugc AB055116, AB055117, AB055118 glutathione S-transferase C. maxima **
CYP88A AF212990, AF212991 cytochrome P450; ent-kaurenoic acid oxidase (multiple alleles) C. maxima ‘Queensland Blue’ 23
PP2 AF520583 phloem protein 2 C. digitata PI 240879 **
PP2 AF520582 phloem lectin C. argyrosperma subsp. sororia **
  L32700, L32701 phloem lectin C. argyrosperma 5
  X56948 malate synthase Cucurbita sp.*** ‘Kurokawa Amakuri Nankin’ 44
pMCPN60 X70867, X70868 chaperonin 60 ‘Kurokawa Amakuri Nankin’ 59
PCPK AY07280, AY072802 phloem calmodulin-like protein kinases C. maxima ‘Big Max’ 66
  X55779 ascorbate oxidase C. maxima ‘Ebisu Nankin’ 14
AAO D55677 ascorbate oxidase C. maxima 33
chitP1 AB015655 chitinase C. maxima ‘Ebisu Nankin’ **
PLC AF082284 chitinase C. moschata crookneck 32
PV72 AB006809 vacuolar sorting receptor ‘Kurokawa Amakuri Nankin’ 54
  D88420 stromal ascorbate peroxidase ‘Kurokawa Amakuri Nankin’ 42
  D78256 isocitrate lyase ‘Kurokawa Amakuri Nankin’ 41
  D70895 3-ketoacyl-CoA thiolase ‘Kurokawa Amakuri Nankin’ 31
  D83656 thylakoid ascorbate peroxidase ‘Kurokawa Amakuri Nankin’ 64
  D49433 hydroxypyruvate reductase ‘Kurokawa Amakuri Nankin’ 21
MP28 D45078 membrane protein ‘Kurokawa Amakuri Nankin’ 28
  D38132 glyoxysomal citrate synthase ‘Kurokawa Amakuri Nankin’ 30
  D29629 aconitase ‘Kurokawa Amakuri Nankin’ 19
  D16560 prepro2S albumin ‘Kurokawa Amakuri Nankin’ 17
  D14044 glycolate oxidase ‘Kurokawa Amakuri Nankin’ 58
  AF002016 acyl CoA oxidase ‘Kurokawa Amakuri Nankin’ 18
PP36 AF274589 cytochrome b5 reductase C. maxima ‘Big Max’ **
pAPX AB070626 peroxisomal ascorbate peroxidase ‘Kurokawa Amakuri Nankin’ 48
CM-ACS3 AB038559 ACC synthase C. maxima 62
CmATS AB049135 acyl-(acyl-carrier protein); acyltransferase C. moschata ‘Shirogikuza’ **
  Y00771 glycerol-3-phosphate acyltransferase transit peptide C. moschata ‘Shirakikuza’ 29
  AB002695 aspartic endopeptidase C. pepo 24
PS-1 AF284038 phloem serpin C. maxima 65
SLW AF170086, AF170087 silverleaf whitefly-induced protein (multiple genes) C. pepo zucchini ‘Chefini’ 60
aprX Y17192 anionic peroxidase C. pepo zucchini ‘Black Beauty’ 6
cpCPK1 U90262 calcium-dependent calmodulin-independent protein kinase C. pepo zucchini 13
PP16 AF079170, AF079171 mRNA movement protein; phloem transport (multiple alleles) C. maxima ‘Big Max’ 63
AOBP D45066 transcription factor binding to ascorbate oxidase C. maxima 34
accW D01032 auxin-induced 1-aminocyclopropane-1-carboxylate synthase C. maxima ‘Ebisu’ 47
  U37774 auxin-induced 1-aminocyclopropane-1-carboxylic acid synthase C. maxima 46
ACC1 M58323 1-aminocyclopropane-1-carboxylate synthase C. pepo 52
ACC1A, ACC1B M61195 1-aminocyclopropane-1-carboxylate synthase (2 genes, tightly linked) C. pepo zucchini 26
PHP-1 D86306 proton-translocating inorganic pyrophosphatase C. moschata crookneck **
PP1 U66277 phloem filament protein C. maxima ‘Big Max’ 9
pfiAF4 X81647 trypsin inhibitor C. maxima ‘Supermarket Hybrid’ 45
pfiBM7 X81447 chymotrypsin inhibitor C. maxima ‘Supermarket Hybrid’ 45
  M15265 phytochrome C. pepo zucchini ‘Black Beauty’ 53
NADH M33154 nitrate reductase C. maxima 11
  M36407 11S globulin beta-subunit ‘Kurokawa Amakuri Nankin’ 20
  AF206895 18S ribosomal RNA C. pepo **
  AF479108 26S ribosomal RNA C. pepo 56
  AJ488214 EF595858 FJ915115 FJ915114 FJ915113 FJ915112 FJ915111 FJ915110 FJ915109 FJ915108 FJ915107  FJ915106 FJ915105 FJ915104 FJ915101 AM981172 AM981170 AM981169 AM981168 5.8S ribosomal RNA C. moschata
C. ficifolia
C. pepo
C. lundelliana
**, 7
  AY396415 5S ribosomal RNA C. pepo 12
  FJ263619 16S ribosomal RNA C. moschata **
  DQ298735 AY357209 AY357208 18S ribosomal RNA C. pepo
C. moschata
  AF017158 25S ribosomal RNA C. maxima **
GID1b AM745267 gibberellin receptor C. maxima **
APRX DQ518906 class III peroxidase precursor C. pepo zucchini‘Black Beauty’ 10
RBP50 EU793994 polypyrimidine tract binding protein C. maxima‘Big Max’ 16
  AJ829947 reverse transcriptase C. pepo **
rbcL AF206756 L21938 DQ535804 EU309692 ribulose 1,5-bisphosphate carboxylase C. pepo
C. ficifolia
C. moschata
NACP1 FJ151402 NAC-domain containing protein C. maxima 50
DNCED1 EU391616 9-cis-epoxycarotenoid dioxygenase C. moschata **
PhoH1 AB435244 alpha-1,4-glucan phosphorylase H isozyme C. maxima **
PhoL1 AB435243 alpha-1,4-glucan phosphorylase L isozyme C. maxima **
PP16-1 EU430061 16kDa phloem protein 1 C. maxima´
C. moschata ‘Ribenzhenmu’
PP16-2 EU430062 16kDa phloem protein 2 C. maxima´
C. moschata ‘Ribenzhenmu’
PP16-1 EF055181 phloem protein 1 C. pepo **
PP16-2 EF055182 phloem protein 2 C. pepo **
  D01033 1-aminocyclopropane-1-carboxylate synthase C. maxima ‘Ebisu’ 27
  EF103124 mitochondrial alternative oxidase C. pepo **
matK DQ536666 DQ536665 DQ536664 maturase K C. pepo
C. digitata
C. ficifolia
trnG EF595908 tRNA-Gly C. pepo 15
  EF202177 aquaporin C. ficifolia **
  EU056338 chitinase C. moschata **
cat1 D55645 catalase C. pepo **
cat2 D55646 catalase C. pepo **
cat3 D55647 catalase C. pepo **
  AF260737 catalase C. pepo **
FTL1 EF462211 DQ865290 flowering locus T protein 1 C. moschata PI441726
C. maxima ‘Big Max’
FTL2 DQ865291 flowering locus T protein 2 C. maxima ‘Big Max’ 40
  AB303333 glyoxalase I C. maxima **
  EF062594 Cu-Zn SOD C. ficifolia **
EF199760 EF199759 EF199758 EF199757 EF199756 EF199755
NBS resistance protein C. moschata **
  AB002695 aspartic endopeptidase C. pepo 24
DHAR EF122791 dehydroascorbate reductase C. ficifolia **
API DQ286449 DQ286448 DQ286447 DQ286445 DQ286444 DQ286443 DQ287856 aspartic acid proteinase inhibitor C. pepo
C. maxima
  EF055184 EF055183 EF055180 16 kDa phloem protein 2 C. moschata
C. ficifolia
PP16 DQ088368
16 kDa. phloem protein 2 C. maxima ‘Lefki kolokytha’ **
PATL1 DQ251455 patellin 1 C. pepo ‘Fordhook’ 49
  E02079 glycerol-3-phosphate acyltransferase C. moschata **
  AJ628045 AJ630372 histidine kinase C. maxima **
A215 X76086 14-3-3 protein endonuclease C. pepo 43
EIN3 DQ023224 DQ023223 EIN3-like protein C. moschata **
aprx Y17192 peroxidase C. pepo zucchini ‘Black Beauty’ **
pfiAF4 X81647 fruit trypsin inhibitor C. maxima ‘Supermarket Hybrid’ **
pfiBM7 X81447 chymotrypsin inhibitor C. maxima‘Supermarket Hybrid’ **
  X73314 Gibberellin 20-oxidase C. maxima‘Riesenm Elone, Gelb Genetzt’ **
  X55779 ascorbate oxidase Cucurbita spp. ‘Ebisu Nankin’ **
pMCPN60-2 X70867 X70868  X68606 chaperonin 60 Cucurbita spp. ‘Kurokawa Amakuri’ **
  AJ829946 AJ829945 AJ829944 reverse transcriptase C. pepo **
NIP1 AJ544830 Nod26-like protein C. pepo 35
GAIP-B AY326307 AY326306 gibberellic acid insensitive phloem B C. maxima 22
  AY663852 serine/threonine kinase-like protein C. ficifolia **
CPR AB116239 oxidosqualene cyclase C. pepo **
CPQ AB116238 cucurbitadienol synthase C. pepo **
  AY672635 chymotrypsin protease inhibitor C. maxima **
  AY672634 aspartic protease inhibitor C. maxima **
AIG-2 AY666083 aspartic protease inhibitor C. maxima **
AIG-1 AY666082 aspartic protease inhibitor C. maxima **
rpl2 AY396281 ribosomal protein L2 C. pepo 12
rpl23 AY396396 ribosomal protein L23 C. pepo 12
rps19 AY396376 ribosomal protein S19 C. pepo 12
psbC AY396185 photosystem II protein C. pepo 12
rpoB AY396320 polymerase beta subunit C. pepo 12
rps2 AY396301 ribosomal protein S2 C. pepo 12
FAD2 AY525163 omega-6 fatty acid desaturase C. pepo **
matR AY453101 maturase C. pepo 3
GAS1 AY379783 galactinol synthase C. pepo 2
atpB AF209573 ATP synthase beta subunit C. pepo **
Pugf AB059484 glutathione S-transferase C. maxima 25
nad1 nad2 AF453584
NADH dehydrogenase subunit 1 and 2 C. pepo ssp. pepo
C. pepo ssp. fraterna
C. pepo ssp. ovifera
C. pepo var. texana
C. pepo var. ozarkana
C. moschata
C. maxima
C. foetidissima
C. argyrosperma
C. sororia
C. ecuadorensis
C. andreana;
C. okeechobeensis ssp. martinezii
CmMP73 AB062669 preproMP73 C. maxima‘Kurokawa Amakuri Nankin’ **
CmATS1;2 AB042401 AB042400 glycerol-3-phosphate acyltransferase C. moschata **
  AF260736 glucose-6-phosphate dehydrogenase C. pepo **
  AF260735 AF260734 AF260733 AF260732 NADP-dependent malic enzyme C. pepo **
  AF260731 heat shock protein 70 C. pepo **
API-2 API-1 AF038167 AF038166 aspartic proteinase inhibitor C. maxima 8
* Gene symbols were assigned by the researchers isolating the gene; they have no correspondence to the official Cucurbita gene symbols.
**Unpublished: Genes can be submitted directly to Genbank, wthout being published in a journal.
*** ‘Kurokawa Amakuri Nankin’ was identified only as “Cucurbita sp.”

Literature Cited in Table 4

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