HORTSCIENCE 26(12):1541-1543. 1991. through the selection of horticulturally su- perior seedlings for asexual propagation as Development of Tetraploid Hybrid clonal cultivars. The highest degree of en- hancement envisioned was the development Clones with Potential for of a group of cross-compatible cultivars combining the cold tolerance common in P. incarnata with fruit quality equal to the best the North Temperate Zone now available in improved cultivars of P. R.J. Knight, Jr. edulis. This paper documents the progress to date toward the stated objective. Subtropical Horticulture Research Station, Agricultural Research Service, In Summer 1979, a clone of P. incarnata U.S. Department of Agriculture, Miami, FL 33158-1399 collected by H.F. Winters in Maryland was Additional index words. edulis, P. incarnata, P. incarnata × P. edulis, crossed with three introductions of P. edulis: polyploidy, colchicine, fertility P.I. 424813, a purple-fruited form collected wild in ; P. edulis f. flavicarpa (M- Abstract. When L. was crossed with P. edulis f. flavicarpa De- 17236, yellow-fruited); and an intraspecific gener, all of the diploid hybrid were pollen-sterile and nonfruitful. Doubling the hybrid (M-21471) derived from crossing a chromosome number of emergent FI seedlings with colchicine restored fertility in some purple-fruited strain from , ‘Nor- individuals, but all plants were strongly self-incompatible and many showed low pollen folk Island’, with a clone of P. edulis f. flav- viability. Crossing colchicine-treated plants that had been converted to amphiploids icarpa (P. I. 243804) brought to from produced a tetraploid hybrid group of four seedling progenies that had some degrees Trinidad (Table 1). Seedlings from these of cross-compatibility. of the amphiploid hybrid is lighter in color than that of crosses were treated upon emergence with a P. edulis, but is sweet, strongly aromatic, and may have use alone or, typically,. as a 0.2% aqueous solution of colchicine to dou- blend with other . ble ploidy and thus restore fertility. When the began to flower in Spring 1980, I In 1971, a clone of maypop (Passiflora Because of sterility of such diploid interspe- evaluated pollen fertility and tried to obtain incarnata) collected in Tennessee was crossed cific hybrids (2n = 18; Darlington and Wy- successful because none of the with P. Cincinnati Masters (P.I. 98883), a lie, 1955), an initial and essential aspect of treated plants proved to be self-compatible. passion from with spectacu- the overall objective was to recover fertility Three of the colchicine-treated seedlings were larly colorful flowers. The F, hybrid, named by doubling the chromosome number of F1 successful seed parents and four succeeded ‘Incense’, was released in 1973 (U.S. Dept. hybrids to produce a group of tetraploid plants as pollen parents (Table 1). The seven fertile of Agriculture, 1976) and continues to be capable of crossing among themselves even colchiploids were intercrossed to produce four sold in the nursery trade. ‘Incense’ combines though they could not be backcrossed to either families of seedlings that were used in later the ornamental value of its pollen parent with of the diploid parental because of the work. One of these, M27461 (M26553A × the ability of P. incarnata to survive tem- difference in ploidies. The tetraploid hybrid M26654B), along with one of the original perate-zone winters in the vicinity of Wash- group had to have a genetic base sufficiently colchiploids, M26509A, was used to pro- ington, D.C. (Winters and Knight, 1975). broad to permit germplasm enhancement duce the progenies that were evaluated in As with its seed parent, all aboveground parts of ‘Incense’ die in the first freezing weather of late autumn, but the vine resumes growth in spring and begins to flower in early sum- mer. Some interest exists now in producing passion fruit in the temperate zone as an an- nual crop (Dozier et al., 1991). The survival of ‘Incense’ for several years in the Wash- ington, D, C., area suggested the possibility of using P. incarnata as a source of winter hardiness to combine with the fruit quality of P. edulis and P. edulis f. flavicarpa to obtain an edible passion fruit for use as a perennial crop in temperate-zone climates. The hybrid between P. incarnata and P. ed- ulis f. flavicarpa was obtained, but like ‘In- cense’, it proved to be pollen-sterile (Knight, 1974). The objective of the work reported here was to develop a group of fertile plants of interspecific hybrid origin, using P. edulis and P. incarnata as the parents, that would be capable of functioning as perennial fruit crop cultivars in temperate-zone climates to which forms of Passiflora edulis-the purple and yellow passion fruits—are not adapted.

Received for publication 30 May 1991. I thank Guy J. Hallman, research entomologist, for as- sistance with statistical analysis of data. The cost of publishing this paper was defrayed in part by the payment of page charges. Under postal regu- lations, this paper therefore must be hereby marked advertisement solely to indicate this fact.

HORTSCIENCE, VOL. 26(12), DECEMBER 1991 1541 Table 3. Response of tetraploid hybrid Passiflora clones to by clone M30173H and/or M30173H x M30174C resulted in 39 seeds, M30174C. whereas M30174C x M30173H yielded a mean of 28 seeds. Earlier observations that fruit weight and juice content of passion fruit (P. edulis) are directly related to the number of seeds matured are confirmed by the con- sistency of juice per seed in this tetraploid hybrid group. In southern Florida, plants of the tetra- ploid hybrid group are similar to vines of P. edulis f. flavicarpa in that, after a normal winter, they start flowering in spring (from zMean separation by Wailer-Duncan K ratio t test, P = 0.05. late April to May) and cease in late autumn NSNonsignificant. (November or December). Thus, fruit rip- ening is scattered from summer to early win- Table 4. Fruit characters of 10 tetraploid passion fruit clones. ter. Characteristics of 10 clones from the te- traploid hybrid group were compared for specific attributes. color of ripe fruit varied from deep green (M30176B) to deep maroon (M30173J and all M30174 seedlings reported here) and, in some cases, was mot- tled-either dark red on a brown background (30173H) or dark red on a green background (all M30177 seedlings reported) (Table 4). Fruit weight varied from 0.5 to 97.8. The number of seeds per fruit in the plants ob- served ranged from one to 129. The amount of juice per fruit ranged from 0.5 to 31.5 ml. Total acidity of juice (expressed in grams per liter) ranged from 2.0 for M30174N to 4.4 for M301740. Total soluble solids content for the tetraploid ranged from 13.2% to 17.2%. The initial observations of the tetraploid hybrid group suggest that the interspecific hybrids have promise as a new fruit crop for warm temperate-zone climates. Vines in field plantings at Miami and at the Southern Fruit and Tree Nut Research Lab., Byron, have zG = green, M = maroon, R = red, T = mottled. survived temperatures below freezing, and field evaluations are continuing. External appearance of the solid maroon fruit has def- 1985 and 1986 for horticultural characters brids observed in 1985 and 1986 were strongly inite eye appeal. The peel has a velvety ex- and responses to pollination (Table 2). In- self-incompatible, indicating that a popula- ternal texture that distinguishes it from the dividual fruit were gathered as they ripened, tion of dependable pollinating insects will be waxy surface of fruit of the purple passion and juice was extracted by hand, using needed wherever this crop is grown. Only a fruit. Internal fruit color and juice color is cheesecloth, and stored in screw-top glass minority of the seedlings sampled were suc- lighter than that of P. edulis, a creamy-yel- vials at -20C until all samples could be cessful when their pollen was used in cross- low suggestive of pulp and juice, analyzed. Root tips of 16 newly emerged pollinations, although all those that showed unlike the intense orange-yellow character- seedlings of M27461 were fixed in 3 acetic normal pollen were tried (Table 2). Means istics of P. edulis juice. As passion fruit juice : 1 alcohol (v/v), stained with acetocarmine, for fruit weight, number of seeds per fruit, is often blended with other juices in fruit and their chromosomes were counted in ac- juice per fruit, and juice per seed were sep- drinks, its color is not of critical importance. cordance with standard laboratory proce- arated by analysis of variance using the Dun- Sufficient juice has not been available to per- dure. Fifteen of these consistently showed a can-Waller multiple range test (SAS Institute, mit extensive taste tests to compare accept- somatic number of 36, whereas one, 1985) for eight male-female combinations ability of members of the tetraploid hybrid M264611, had cells that varied in chromo- involving the two most successful polliniz- group with currently available cultivars of P. some number from 34 to 36. Thus, a func- ers, M30173H and M30174C. The degree of edulis. However, based on my subjective tioning tetraploid population had been fertility of various parental combinations in- observations, the juice was pleasant in fla- achieved. volving these two pollinizers, as demon- vor, both acidic and sweet, as the analytical Efforts to use the seedlings of progeny strated by the number of seeds per fruit data indicate (Table 4), and also was strongly M27461 in breeding began in Spring 1981 obtained, differed significantly (Table 3). One aromatic, resembling juice of P. edulis in and continued through Summer 1983. At that seed parent, M30176B, was highly success- this respect. Thus, there is reason to expect time, accession numbers were assigned to 89 ful with both pollen parents. M30176B yielded that juice of selections from the tetraploid seeds from which the seedling clones ex- 90 seeds or more from either pollination and hybrid group can be put to many of the same amined in 1985 and 1986 were derived. One averaged 19.4 and 25.3 ml of juice/fruit, uses as is juice of the purple or yellow pas- exceptional population, M30177, consisted suggesting that, with adequate insect polli- sion fruit. of 17 seedlings resulting from a single self- nation, satisfactory yields can be expected in The self-incompatibility of the hybrid plants pollination of M21471H, the only time self- the field. When the two pollen parents were puts potential growers at a definite disad- pollination of a tetraploid passion fruit hy- used in reciprocal cross-pollinations, both set vantage because it necessitates taking mea- brid succeeded. fruit but neither demonstrated a high degree sures to ensure cross-pollination. Finding self- Without exception, all the amphiploid hy- of cross-compatibility; the combination of compatible plants in seedling populations is

1542 HORTSCIENCE, VOL. 26(12), DECEMBER 1991 necessary. Cultivated forms of P. edulis are Literature Cited fruit production. Proc. XIX Intl. Hort. Congr. (Warsaw) 3:59-68. known to encompass self-compatible geno- Darlington, C.D. and A.P. Wylie. 1955. Chro- types (Knight and Winters, 1963), but no mosome atlas of flowering plants. 2nd ed. George Knight, R.J. and H.F. Winters. 1963. Effects of self-compatible forms of P. incarnata have Allen and Unwin, London. selfing and crossing in the yellow passion fruit. been confirmed to date. The maypop, how- Dozier, W. A., Jr., R. Rodriguez-Kabana, A.W. Proc. Fla. State Hort. Soc. 76:345-347. ever, has a wide distribution in the southern Caylor, D.G. Himelrick, N.R. McDaniel, and SAS Institute, 1985. SAS/STAT guide for per- and central , ranging from Flor- J.A. McGuire. 1991. Ethephon hastens matu- sonal computers, version 6 ed. SAS Institute, ida and Texas northward to Virginia, Illi- rity of passionfruit grown as an annual in a tem- Inc., Cary, N.C. nois, and southeastern Kansas (Killip, 1960), perate zone. HortScience 26:146-147. and possesses considerable variability. Killip, E.P. 1960. Supplemental notes on the U.S. Dept. of Agriculture. 1976. inventory Therefore, if self-compatible forms of this American species of with de- no. 182. U.S. Dept. Agr. Washington, D.C. species can be found, it would appear desir- scriptions of new species. Contrib. U.S. Natl. Winters, H.F. and R.J. Knight, Jr. 1975. Select- able to use such material in resynthesizing Museum 35(1):1-23. ing and breeding hardy passionflowers. Amer. the interspecific hybrid. Knight, R.J. 1974. Special problems in tropical Hort. 54:22-27.

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