Russet is a disorder of apple (Malus×domestica) fruit where the formation of cork cells leads to a cosmetic blemish which is commercially undesirable. One of the many causes of russet is low temperature damage early in fruit development. Following frost damage to fruit, a study was initiated to determine whether carbaryl chemical thinner was more effective in thinning russeted fruit than nonrusseted fruit. With no chemical thinner application, russeted fruit abscised at a greater rate than nonrusseted fruit. Following the application of carbaryl to the fruit however, there was no difference in the retention of fruit among the treatments. Chemically thinning with carbaryl therefore is not a technique that growers could use to preferentially thin russeted fruit.
Anish Malladi and Peter M. Hirst
Peach production is significantly reduced and severely limited by frost injury in regions frequently exposed to late spring freeze conditions. Peach flower buds become increasingly susceptible to low-temperature damage from the period of completion of rest through fruit set. Delaying dehardening and/or flower bud development is an effective way to avoid frost damage. Bio-regulator applications, affecting dormancy or bud development, can delay flowering and dehardening of the buds and can help in avoiding spring freeze injury. Spring applications of AVG and dormant oils on 8-year-old `Redhaven' peach trees were evaluated. AVG applications effectively delayed bloom by 2 to 5 days. The most effective treatment was two applications of 2000 ppm AVG, which delayed bloom by almost 5 days. Repeat applications of AVG were more effective than the single dosage treatments. The 1000 ppm, repeat application delayed bloom by 4 days. A single application of 5000 ppm AVG resulted in severe phytotoxicity. The wetting agent levels were also varied and AVG applications were most effective in combination with 0.2% `Sylgard'. AVG, apparently, delayed bloom by delaying bud development following the completion of rest. The dormant oil sprays were ineffective in achieving bloom delay. The specific leaf weight characteristics of the treated trees were not affected except for the 5000-ppm AVG application, which reduced SLW. Fruit characteristics such as maturity, weight, and soluble sugar concentration were not affected by any of the spring applications (except for the 5000-ppm AVG application, which was phytotoxic). Our studies indicate that AVG is effective in delaying bloom in peaches by up to 5 days. This has the potential to substantially increase peach yields in years with a late spring freeze.
Peter M. Hirst and Wendy M. Cashmore
Spurs were collected periodically throughout three growing seasons from the 1-year-old section of wood of `Royal Gala' trees growing in New Zealand. Three classes of spurs were sampled: purely vegetative spurs, those that flowered but did not carry fruit, and spurs on which a single fruit was borne. The bourse bud, in which flowers may form for the following year's crop, was dissected and bud appendages classified and counted. In addition, axillary buds from current-season shoots were sampled and dissected. Over the period 50–200 days after full bloom, the number of appendages in buds on vegetative spurs increased from ≈14 to 22, whereas the increase in buds on fruiting spurs was 14 to 20. In contrast, axillary bud appendage numbers increased from ≈11 to 14 over this period. By the end of the growing season, flowers were evident in a high proportion of buds of all classes. The critical appendage number at which the change from a vegetative to floral status became visible was ≈18 for spurs on 1-year-old wood, but 13 for axillary buds. The time at which flowers were able to form varied among years. The degree of flower differentiation that occurred prior to leaf fall was highest in vegetative buds and was reduced by flowering and fruiting, and was lowest in axillary buds.
Peter M. Hirst and David C. Ferree
One way in which rootstocks may influence production efficiency is by altering the number of spurs, and in particular reproductive spurs. However, rootstock influences on the morpholgy of shoots have not been quantified. Measurements were made on `Starkspur Supreme Delicious' trees growing on 17 rootstocks and planted in 1984 as part of the NC-140 regional rootstock trial. In each of the 6 years from 1988-1993, the length of the 2-year old section of wood of selected branches was measured and the number of spurs, flowers and shoots counted. For all rootstocks, trunk cross-sectional area was closely related to shoot length. Trees on P.22 (the most dwarfing rootstock in the planting) had shoot lengths 40-50% of those of trees on seedling rootstocks. For each rootstock, there was a strong negative relationship between shoot length and spur density, but there was not a common relationship among rootstocks. Similarly, flower number per shoot was also related to shoot length with different relationships for each rootstock. Flower density was not related to vigor for any of the rootstocks.
Khalil R. Jahed and Peter M. Hirst
Pollination is an essential prerequisite for the production of many fruit and seed crops, including apple. In apple, successful fertilization requires pollen transfer to the stigma, pollen germination, and successful pollen tube growth resulting in fruit set. Precise selection of the most effective pollinizers for commercial orchards is not possible however, until these processes are more fully understood. The present study was undertaken to compare pollinizers in terms of pollen tube growth and fruit set. On trees of ‘Honeycrisp’, ‘Fuji’, and ‘Gala’ from which bees were excluded, flowers were hand-pollinated using pollen collected from crabapple (‘Ralph Shay’ or Malus floribunda), ‘Delicious’ and ‘Golden Delicious’. Flowers were harvested at one, two, three, and four days after pollination (DAP). Pollen source had a significant influence on pollen germination on the stigmatic surface, number of pollen tubes penetrating the stigma, distance of pollen tube growth down the style, and pollen tubes reaching the base of the style. In ‘Honeycrisp’ and ‘Gala’, ‘Golden Delicious’ pollen grew the fastest, followed by ‘Delicious’ and crabapple. Neither ‘Ralph Shay’ nor Malus floribunda were effective pollinizers for ‘Honeycrisp’ and resulted in low fruit set suggesting incompatibility may be involved. However, both these crabapples were effective pollinizers for ‘Fuji’ and ‘Gala’. These results indicate that pollen source can have a tremendous impact on pollen tube growth and fruit set. The physiological basis for these effects is not clear, but implications for pollinizer selection are obvious.
Mokhles A. Elsysy and Peter M. Hirst
Lack of consistent flower formation is the underlying cause of biennial bearing. Flower formation in apple (Malus ×domestica Borkh.) has been associated with different factors, such as leaf area, shoot growth, bourse length, crop load, and seed number. However, it is unclear how these different factors interact to promote or inhibit flower formation. The effect of spur defoliation, fruit removal, and their interaction were evaluated on spur flower formation and bourse length in annual-bearing ‘Gala’ and the biennial-bearing ‘Honeycrisp’. Eight different combinations of spur defoliation and fruiting treatments were applied in three consecutive springs, 2013–15. Bourse shoot defoliation and fruiting treatments inhibited spur flower formation in both cultivars, but in different patterns from year to year. In addition, spur leaf defoliation did not affect flower formation in either cultivar. Furthermore, local defoliation and fruiting treatments did not affect bourse length. We propose that bourse leaves play a major role in both producing and transporting flower formation signals, but the effect depends on cultivar.
Peter M. Hirst and David C. Ferree
Two-year-old branch sections of `Starkspur Supreme Delicious' apple (Malus domestics Borkh.) trees growing on 17 rootstock were studied over 6 years to determine the effects of rootstock on shoot morphology and spur quality and describe how these factors may be related to precocity and productivity. Shoot length was affected by rootstock and was positively related to trunk cross-sectional area within each year, but the slope of the regression line decreased as trees matured. The number of spurs on a shoot was largely a product of shoot length. Spur density was inversely related to shoot length, where rootstock with longer shoots had lower spur densities. Flower density was not related to spur density, and shoot length only accounted for a minor part of the variation in flower density. The proportion of spurs that produced flowers was closely related to flower density, indicating that rootstock influence flower density by affecting the development of individual buds rather than by the production of more buds. More vigorous rootstock generally had spurs with larger individual leaves and higher total leaf area per spur, but fewer spur leaves with lower specific leaf weights. More precocious rootstock were also more productive over a 10-year period when yields were standardized for tree size. Tree size was the best indicator of precocity and productivity, which could be predicted with a high degree of certainty as early as the 4th year.
Peter M Hirst and David C Ferree
Floral development was studied in buds of `Starkspur Supreme Delicious' apple trees growing on B.9, M.26 EMLA, M.7 EMLA, P.18, and seedling rootstocks. In each of 3 years, buds were sampled from the previous years growth at intervals throughout the growing season and dissected to determine whether the apex was domed, indicating the start of floral development. Number of bud scales and true leaves increased during the early part of the growing season, but remained fairly constant beyond 70 days after full bloom. The type of rootstock did not affect the number of bud scales or transition leaves, and effects on true leaf numbers were small and inconsistent. Final bract number per floral bud was similarly unaffected by rootstock. The proportion of buds in which flowers were formed was influenced by rootstock in only one year of the study, which was characterized by high temperatures and low rainfall over the period of flower formation. Bracts were observed only in floral buds, and became visible after doming of bud apices had occurred. Flowers were formed during the first 20 days in August, regardless of rootstock or year. The appendage number of vegetative buds was constant from 70 days after full bloom until the end of the growing season, but the number of appendages in floral buds increased due to the continued production of bracts. The critical bud appendage number for `Starkspur Supreme Delicious' before flower formation was 20, and was stable among rootstocks and years. Buds with diameters above 3.1 mm were generally floral, but on this basis only 65% of buds could be correctly classified. Spur leaf number, spur leaf area, and spur leaf dry weight were not good predictors of floral formation within the spur bud.
Peter M. Hirst and David C. Ferree
In each of 3 years, vegetative spurs were sampled from l-year-old wood of `Starkspur Supreme Delicious' apple trees (Malus domestica Borkh.) growing on B.9, M.26 EMLA, M.7 EMLA, P.18, and seedling rootstocks. Mineral concentrations of spur leaves and bud apical meristems were determined, and related to spur bud development. The spur leaf P concentration decreased during the growing season each year, hut was unaffected by rootstock. Spur leaves of trees on B.9 rootstock had 30% higher Ca concentrations than trees on M.26 EMLA or seedling rootstocks. In each year, trees growing on M.26 EMLA rootstocks had the highest leaf Mg concentrations. Mineral concentrations were generally unrelated to spur leaf number, leaf area, leaf dry weight, or specific leaf weight. Phosphorus concentrations in spur bud apical meristems declined during two of the three growing seasons of the study and were unaffected by rootstock. Bud P concentration was weakly negatively related to bud diameter and bud appendage number in one year of the study. More vigorous spurs (as indicated by higher spur leaf number, leaf area, and leaf dry weight) had higher bud K levels during each year. No relationships between bud development and either spur leaf mineral concentration or bud apical meristem mineral levels were evident, suggesting that a direct role of mineral nutrition influenced by rootstock at the site of flower formation was unlikely.
Mokhles A. Elsysy and Peter M. Hirst
Adequate flower formation limits dependable apple (Malus ×domestica) production and is a major challenge for apple industries around the world. ‘Honeycrisp’ is a high value apple cultivar, but consistent flowering is difficult to achieve. Apple flower formation is affected by factors including defoliation, girdling, and gibberellin (GA4+7) and 1-naphthaleneacetic acid (NAA) applications. However, the molecular mechanisms that regulate the effects of these factors are not well understood. We studied the effect of local spur defoliation, GA4+7 and NAA applications on ‘Honeycrisp’ flower formation. Furthermore, we investigated the effect of local defoliation and local GA4+7 application on the transcript levels of two major flower formation genes in the meristems of apple spurs. The floral inhibition gene terminal flower1-1 (MdTFL1-1) and floral promoting genes flowering locus T (MdFT1 and MdFT 2) of apple. Local application of GA4+7 and defoliation treatments inhibited flower formation, but NAA applications were without effect. Defoliation treatments were accompanied by a significant reduction in MdFT1, 2 transcript levels compared with controls early in the growing season. Conversely, GA4+7 application was accompanied by a significant increase in MdTFL1-1 transcripts compared with controls throughout the growing season. These results indicate that GA4+7 inhibits flower formation by upregulating the inhibitory MdTFL1-1, and defoliation acts by downregulating transcript levels of MdFT1, 2 early in the growing season. We also provide evidence that defoliated bourse buds may receive flowering promotion signals from other parts of the tree in the absence of their local leaves.