The temperature and ethylene response of ripening papaya fruit (Carica papaya L. cv. Sunset) was determined with and without 14 days of storage at 10C. Temperatures at or higher than 30C adversely affected the quality of the ripe papaya. Papayas held at 32.5C for 10 days failed to ripen normally, as evidenced by poor color development, abnormal softening, surface pitting, and an occasional off-flavor. Skin yellowing, fruit softening, and flesh color of papayas exhibited a quadratic response to ripening time within the temperature range of 22.5 to 27.5C. Flesh color development of nonstored fruit did not change significantly during the first 6 days at ripening temperatures, then rapidly increased. Fruit stored for 14 days at 10C exhibited faster ripening rates (e.g., degreening and softening and no delay in flesh color development) than nonstored fruit when removed to other ripening temperatures (17.5 to 32.5 C). Problems of weight loss and development of external abnormalities were more significant at temperatures higher than 27.5C. The optimal temperature range was found to be between 22.5 and 27.5C, with fruit taking 10 to 18 days to reach full skin yellowing from color break, whether or not fruit was stored at 10C. Exogenously applied ethylene (=100 μl·liter-1) stimulated the rate of fruit ripening, as measured by more uniform skin yellowing and rate of flesh softening whether or not the fruit were stored for 14 days at 10C. Ethylene did not ripen immature papayas completely in terms of skin and flesh color development. The outer portion of the flesh of ethylene-treated fruit had a faster rate of ripening, as indicated by carotenoid development and softening rate, while the same area of the flesh was still pale white in nonethylene-treated fruit. Ethylene reduced the coefficient of variation for skin color, softening rate, and flesh color development in treated fruit. Ethylene increased the rate of skin degreening and hastened the rate of carotenoid development and softening in the outer mesocarp, while having little effect on the inner mesocarp.
Nine transgenic papaya clones, produced previously by microprojectile bombardment, are being characterized for frequency of somaclonal variation. Five clones have proven to be hermaphrodite. Four of these appear to have normal fertility, while the fifth has drastically reduced pollen fertility, averaging about 15% stainability with acetocarmine. Four other clones are pistillate and appear to have normal fertility, with one exception which has been demonstrated to be tetraploid (2n=36 chromosomes). One of twelve plants in a pistillate clone was a somaclonal mutant showing altered leaf and flower morphology. The transgenic clones and their sexual progenies are also being evaluated at the molecular level for expression and segregation of npt, gus, and the coat protein (CP) of papaya ringspot virus (PRV), as well as for PRV resistance.
Gynodioecious papaya (Carica papaya L.) seedlings in commercial cropping systems in Hawaii are typically multiple-planted and thinned upon flowering to a single hermaphrodite because seedlings segregate for sex expression. Use of clonally propagated hermaphrodites would eliminate the over-planting practice and may provide other advantages. Yields of clonally propagated hermaphrodites were compared with single- and multiple-planted seedlings in three fields on two islands in Hawaii. Cloned hermaphrodites were either rooted cuttings or in vitro micropropagated plants. Clonally propagated plants bore ripe fruit 1 to 3 months earlier than thinned seedlings and had significantly higher early and cumulative yields. At each site, cumulative yields of thinned seedlings never reached the same level as those of clonally propagated plants. The yield benefit from clonally propagated plants was greatest at Keaau, the lowest sunlight and least productive test site.
The occurrence of intra-ovarian ovaries in certain strains of Carica papaya L. is not uncommon. Approximately 150 hermaphroditic and 25 female ovaries from a hybrid progeny contained internal ovaries in stages of development ranging from thread-like appendages to round or elongated pistils of various sizes and shapes. A few were large enough to fill the entire seed cavity of the primary fruit and possessed their own cavities with non-viable seeds.
Internal ovaries originate either from stimulated growth of rudimentary pistillate structures extending from the central axis of the receptacle or from placentae in positions normally occupied by ovules. The placenta may be in its normal parietal position or a single strand may become free, extended from the base of the primary ovary and support a mal-shaped secondary ovary as well as ovules. The occurrence of internal ovaries supports theories proposed by other investigators on the evolution of floral morphology in the papaya.
Genetically engineered (GE), virus-resistant papaya cultivars in Hawaii are easily identified by a colorimetric assay for the β-glucuronidase (GUS) marker transgene. We used GUS to track pollen movement from a central 1-acre plot of gynodioecious GE `Rainbow' plants into seeds on surrounding border rows of non-GE `Sunrise' papaya. GUS evidence of cross-pollination occurred in 70% of female plants (43% of assayed seeds), compared with only 13% of the predominantly self-pollinating hermaphrodite plants (7% of seeds) segregating in the gynodioecious `Sunrise' border rows. The percentage of GUS+ seeds in border row plants showed a weak negative correlation (r = –0.32) with distance from the nearest GE tree (30 m maximum). In a non-GE papaya field located less than a mile downwind from the `Rainbow' source, no evidence of GUS was found in 1000 assayed seeds. In a separate study, the origin of GUS+ seed discovered in papaya fruits from an organic farm was investigated. Leaf GUS assays revealed that 70% of trees were GE, indicating that the grower had planted GE seed. The impact of pollen drift from GE trees in the same field was determined by screening seed samples from 20 non-GE hermaphrodites for GUS expression. Only three hermaphrodites (15%) showed GUS+ seeds, at low levels ranging from 3% to 6% of contaminated samples. These data indicate that the major source of GE contamination in organic fields is seeds of unverified origin, rather than pollen drift from neighboring GE fields. Organic growers are advised to: 1) plant only seed that is known to be non-GE, preferably obtained by manual self-pollination of selected non-GE hermaphrodites; 2) avoid open-pollinated seed; and 3) grow only hermaphrodite (self-pollinating) trees, removing any female or male plants from production fields.
`Solo' papaya (Carica papaya L.) fruit removed at different points from a commercial packing house showed that skin injury due to mechanical damage increased as fruit moved through the handling system. The occurrence of “green islands” -areas of skin that remain green and sunken when the fruit was fully ripe-apparently were induced by mechanical injury. Skin injury was seen in fruit samples in contact with the sides of field bins, but not in fruit taken from the center of the bins. Bruise-free fruit at different stages of ripeness (5% to 50% yellow) were dropped from heights of 0 to 100 cm onto a smooth steel plate to simulate drops and injury incurred during commercial handling. No skin injury occurred, although riper fruit showed internal injury when dropped from higher than 75 cm. Fruit (10% to 15% yellow) dropped onto sandpaper from a height of 10 cm had skin injury symptoms similar to those seen on fruit from the commercial handling system. These results suggest that abrasion and puncture injury were more important than impact injury for papaya fruit. Heating fruit at 48C for ≈6 hours or until fruit core temperature (FCT) reached 47.5C aggravated the severity of skin injury. Delays in the application of heat treatment from dropping did not reduce the severity of skin injury significantly, except for fruit heated 24 hours after dropping. Waxing fruit alleviated the severity of skin injury, whether applied before or after the heat treatment. Skin injury to papaya was caused by abrasion and puncture damage-not impact-and increased during postharvest handling of the fruit. The injury was associated mainly with fruit hitting the walls of wooden bins-bin liners may reduce this injury.
Municipal solid waste compost was applied with a side delivery applicator on top of the bed as a mulch in May 1993, 6 months after transplanting at Homestead, Fla. Papaya (`Know You No 1') was grown with and without compost mulch. Compost was distributed on the surface of the bed ≈90 cm wide and 5 cm thick. There were no mulch effects on trunk diameter nor plant height. Plant height was affected by papaya sex 4 and 6 months after transplanting. Hermaphroditic plants were taller than female plants. There were no mulch effects on marketable yield per plant, marketable size, or number of cull fruit. Sex, however, influenced papaya size and total cull number. Hermaphroditic plants produced larger marketable fruit and more cull fruits than female plants. Lower plant mortality rates were found after 1.5 years in the mulched plants compared to unmulched plants. Soil and tissue analysis showed no differences in N, P, K, Mg, S, Mn, Fe, Cu, and B, except for Zn. Zinc contents in soil and tissue were higher in the mulched areas than unmulched areas.
Mesocarp softening during papaya (Carica papaya L.) ripening was impaired by heating at 42C for 30 min followed by 49C for 70 min, with areas of the flesh failing to soften. Disruption of the softening process varied with stage of ripeness and harvest date. The respiratory climacteric and ethylene production were higher and occurred 2 days sooner in the injured fruit than in the noninjured fruit that had been exposed to 49C for only 30 min. Skin degreening and internal carotenoid synthesis were unaffected by the heat treatments. Exposure of ripening fruit to either 42C for 4 hr or 38 to 42C for 1 hr followed by 3 hr at 22C resulted in the development of thermotolerance to exposure to the otherwise injurious heat treatment of 49C for 70 min. Four stainable polypeptide bands increased and seven declined in single-dimensional acrylamide gels following incubation of fruit at the nondamaging temperature of 38C for 2 hr. Three polypeptides showed marked increases when polysomal RNA was translated. These polypeptides had apparent molecular weights of 17, 18, and 70 kDa. Proteins with molecular weights of 46, 54, and 63 kDa had slight increases after heat treatment. The levels of these polypeptides peaked 2 hr after heat treatment and declined within 24 hr. The amount of these polypeptides in the unheated control varied with the batch of fruit. The concentration of three translated polypeptides, with apparent molecular weights of 26, 37, and 46 kDa, declined. Other polypeptides continued to be translated during and after holding papayas for 2 hr at 38C.
Transgenic papaya line 55-1 with resistance to papaya ringspot virus (PRSV) originated in 1989 by particle bombardment of cultivar Sunset with the coat protein gene (cp) of mild mutant Hawaii PRSV strain HA 5-1. Hemizygous (+/cp) R0 clones of 55-1 displayed resistance to the virulent Hawaii HA strain in greenhouse tests in New York in 1991 and to local strains in a field trial in Hawaii from 1992 to 1994. In the R1 generation produced by crossing the pistillate R0 55-1 with `Sunset', up to 50% of the hemizygous transgenic segregants were susceptible to a local Oahu PRSV strain when inoculated as seedlings but not as mature plants. Similar inoculation experiments in New York showed that hemizygous R1 transgenics were susceptible in differing degrees to PRSV strains from regions other than Hawaii. Homozygous (cp/cp) R2, R3, and R4 populations planted in various locations in Hawaii since 1994 have consistently demonstrated high-level resistance to local strains at all stages of development. When inoculated in New York with eight non-Hawaii PRSV strains, homozygous R3 seedlings were resistant to all but a Thai strain. Transgenic resistance is the result of a complex interaction involving the stage of plant development, transgene dosage, the degree of homology between transgene and challenge virus, and environmental variables. Papaya plants transformed with nontranslatable versions of various cp genes are also highly resistant to PRSV, indicating that the resistance mechanism operates at the RNA level. No loss of resistance due to the appearance of resistance-breaking virus strains or to transgene inactivation has been noted thus far.
Two studies were conducted to determine the influence of pH on papaya seed germination and seedling emergence. The germination test was conducted with `Waimanalo' and `Tainung 1' seeds, using a double layer of filter paper disks in plastic petri dishes placed within a growth chamber. Each dish received 40 seeds, and germination was defined as when the radicle was visible. Disks were wetted daily with nutrient solution adjusted to pH of 3, 4, 5, 6, 7, 8, or 9. Germination began on day 5, and the study was terminated on day 23. Solution pH did not influence germination rate or ultimate germination percentage. `Waimanalo' exhibited 58% germination and `Tainung 1' exhibited 64% germination in this test. The seedling emergence study was conducted with `Waimanalo' seeds using sand culture within a growth chamber. Thirty seeds were planted in 10-cm containers, and the sand was irrigated daily with the solutions from the first study. Emergence was defined as when the hypocotyl hook was visible above the sand. Emergence began on day 10, and the study was terminated on day 30. Solution pH did not influence seedling emergence, and mean emergence was 69% in this study. The results indicate that the seed germination and seedling emergence stages of papaya seedling growth are adapted to a wide range of substrate pH.