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The rootstock plays a large role in modern citrus production because of its influence on tolerance to adverse abiotic and biotic soil-borne stresses, and on the general horticultural characteristics of the grafted scion. In recent years, rootstock has received increased attention as a management strategy to alleviate the devastating effects of the bacterial disease huanglongbing (HLB), also known as “citrus greening.” In commercial citrus nursery production, rootstocks are typically propagated by seed. Because of the increased demand for HLB-tolerant rootstocks, seed supply is often inadequate for the most popular cultivars. Cuttings and tissue culture (TC) propagation are alternative methods to supply adequate quantities of genetically identical rootstocks to be used as liners for grafting. However, there are concerns among nursery owners and citrus growers regarding the possible inferiority of rootstocks that are not propagated by seed. This study investigates the influence of rootstock propagation method on traits of sweet orange trees grafted on four commercially important rootstock cultivars during the nursery stage and during the first year of growth in a commercial citrus orchard. Several of the measured traits during the nursery stage, including rootstock sprouting, grafted tree growth, and root mass distribution were significantly influenced by the rootstock propagation method, but traits were also influenced by the rootstock cultivar. Our results also suggest that for tissue culture-propagated plants, differences in the starting material and the culturing method can affect the grafted tree behavior. Except for canopy spread and scion to rootstock trunk diameter ratio, tree growth during the orchard stage was determined by the combination of propagation method and rootstock, rather than by propagation method alone.
Six new hybrid rootstocks, ‘US-1279’, ‘US-1281’, ‘US-1282’, ‘US-1283’, ‘US-1284’, and ‘US-1516’, were released from the U.S. Department of Agriculture (USDA) citrus breeding program to provide improved tree tolerance to huanglongbing (HLB), the most destructive disease facing the citrus industry in the United States and many other parts of the world. Five of these new rootstocks were released based on field performance in trials with the rootstocks propagated by stem cuttings, rather than the traditional propagation using nucellar seedlings. In this study, we evaluated the fruit, seed, and seedling characteristics of these new rootstocks, along with four other USDA rootstocks of commercial importance. The study included a determination of the percentage of true-to-type and off-type seedlings by both plant morphology and simple sequence repeat (SSR) markers. All 10 rootstocks produced an acceptable number of seeds and good seedling emergence from those seeds. The rootstocks ‘Swingle’, ‘US-802’, ‘US-812’, ‘US-1283’, ‘US-1284’, and ‘US-1516’ had a high percentage of true-to-type seedlings and correspondingly good potential to be propagated by seeds. However, no true-to-type plants were observed among seedlings from the rootstocks ‘US-1279’, ‘US-1281’, and ‘US-1282’, indicating that economical seed propagation will be impossible for these cultivars. The 10 SSR marker sets used in this study were observed to easily differentiate the 10 rootstocks studied, and readily distinguished true-to-type and off-type seedlings among progeny from all 10 rootstock clones. This study presents information of significant value for commercial nurseries involved in propagation of citrus rootstocks, and those involved in citrus rootstock breeding and development around the world. We propose the use of these 10 SSR marker sets as readily applicable for accurate identification of most citrus rootstock cultivars and their true-to-type seedlings.
Candidatus Liberibacter asiaticus (Las) is a phloem-limited bacterium associated with huanglongbing (HLB), one of the most destructive diseases of citrus in Florida and other citrus-producing countries. Natural transmission of Las occurs by the psyllid vector Diaphorina citri, but transmission can also occur through grafting with diseased budwood. As a result of the difficulty of maintaining Las in culture, screening of citrus germplasm for HLB resistance often relies on graft inoculation as the mode of pathogen transmission. This study evaluates transmission efficiencies and HLB progression in graft-inoculated and psyllid-inoculated citrus under greenhouse and natural conditions in the field. Frequencies of transmission in graft-inoculated greenhouse-grown plants varied between experiments and were as high as 90% in susceptible sweet orange plants 6 to 12 months after inoculation. Transmission frequency in a tolerant Citrus × Poncirus genotype (US-802) was 31% to 75%. In contrast, transmission of Las after controlled psyllid inoculation did not exceed 38% in any of four experiments in this study. Whereas the time from inoculation to detection of Las by polymerase chain reaction (PCR) was faster in psyllid-inoculated US-802 plants compared with graft-inoculated US-802 plants, it was similar in graft- and psyllid-inoculated sweet orange plants. HLB symptom expression was indistinguishable in graft- and psyllid-inoculated plants but was not always associated with the number of bacteria in affected leaves. The highest number of Las genomes per gram leaf tissue measured in sweet orange plants was one to four × 107 in graft-inoculated plants and one to two × 107 in psyllid-inoculated plants. Highest numbers measured in tolerant US-802 plants were one to three × 106 and two to six × 106, respectively. Compared with artificial inoculation in a greenhouse setting, natural inoculation of field-grown sweet orange trees occurred at a much slower pace, requiring more than 1 year for infection incidence to reach 50% and a minimum of 3 years to reach 100%.
Huanglongbing (HLB) is a devastating disease of citrus that is found in most citrus production areas around the world. The bacterium associated with HLB resides in and damages the phloem, restricting the movement of photosynthates throughout the plant and leading to tree decline. Considerable root loss can be observed in affected trees even when few disease symptoms are visible aboveground. Root traits can substantially influence tree performance and use of superior rootstocks is one strategy to manage tree health and reduce production losses in a disease-endemic environment. Citrus rootstocks are typically propagated by seed, but due to the increased demand for some of the best-performing cultivars, propagation by other methods is being used to overcome seed shortages. In this research, differences in root architecture and root growth of six different rootstocks propagated by seed, cuttings, and tissue culture, and their influence on the grafted ‘Valencia’ (Citrus sinensis) scion were investigated. A field trial was established in southwest Florida in 2017. Trees were evaluated for their performance during the first 2 years after planting and a subset of trees was excavated for detailed analysis of root architectures and biomass distribution. Significant differences among propagation methods were found for the rootstock trunk diameter and the lateral (structural) root length, which were largest in seed-propagated rootstocks. Most of the other horticultural and root architectural traits were not significantly influenced by the rootstock-propagation method; however, many of the measured variables were significantly influenced by the rootstock cultivar regardless of the propagation method. The results showed that rootstocks propagated by cuttings and tissue culture were similar to seed-propagated rootstocks in their influence on the grafted tree during the early years of growth in the field.
The devastating citrus disease huanglongbing (HLB) associated with the phloem-limited bacteria Candidatus Liberibacter asiaticus (CLas) has caused a more than 70% reduction in citrus production since its discovery in Florida in 2005. Most citrus scion cultivars are sensitive to HLB, whereas some cultivars used as rootstocks are tolerant. Using such tolerant rootstocks can help trees to cope better with the disease’s impact. Evaluating rootstock effects on a grafted scion in the field takes many years, but shorter-term evaluation is imperative to aid in rootstock selection for an HLB-endemic production environment. In this study, we investigated grafted healthy and CLas-infected citrus trees under controlled greenhouse conditions. The objectives were to identify traits suitable for assessing grafted tree tolerance in advance of longer-term field studies and aiding in the selection of superior rootstock cultivars. We assessed 10 commercially important rootstocks grafted with ‘Valencia’ sweet orange scion and with known field performance. At 6, 9, 15, and 21 months after graft inoculation (mai), leaf CLas titers were determined and canopy health was evaluated. Plants were destructively sampled at 21 mai to assess plant biomasses and other physiological and horticultural variables. There was little influence of the rootstock cultivar on CLas titers. Surprisingly, few HLB foliar disease symptoms and no differences in soluble and nonsoluble carbohydrate concentrations were measured in infected compared with healthy plants, despite high CLas titers and significant reductions in plant biomasses. Most trees on rootstocks with trifoliate orange parentage were less damaged by HLB than other rootstocks, although results did not always agree with reported field performance. Among the different variables measured, leaf size appeared to be most predictive for grafted tree assessment of HLB sensitivity. The results of this study provide a better understanding of the strengths and weaknesses of assessing rootstock influence on grafted tree performance in a controlled greenhouse environment. Although such studies provide valuable information for cultivar tolerance to HLB, other rootstock traits will ultimately contribute to field survival and productivity in an HLB endemic production environment.
Grafting a scion onto a rootstock results in physical and physiological changes in plant growth and development, which can affect tree vigor, productivity, and tolerance to stress and disease. Huanglongbing (HLB) is one of the most destructive citrus diseases and has become endemic in Florida since its introduction in 2005. It is associated with the phloem-limited bacteria Candidatus Liberibacter asiaticus (CLas), which cause severe metabolic disruptions in affected plants. Although most scion cultivars are highly susceptible, some rootstock cultivars are tolerant and allow the grafted tree to cope better with the disease. The objectives of this study were to identify rootstock traits that can be used to assess cultivars under controlled greenhouse conditions in advance of longer-term field trials. We used 10 commercially important rootstocks with different genetic backgrounds and known field performance in graft combination with ‘Valencia’ sweet orange scion. Trees were graft-inoculated with CLas and compared against mock-inoculated trees. Tree health and CLas populations were assessed regularly, and root growth was monitored using a minirhizotron imaging system. Plants were excavated and destructively sampled 21 months after inoculation to assess biomass distributions and other CLas-induced effects. We found significant differences between healthy and infected trees for most variables measured, regardless of the rootstock. In contrast to leaf CLas titers, root titers were significantly influenced by the rootstock, and highest levels were measured for ‘Ridge’ sweet orange and sour orange. Root growth and root biomasses were reduced upon infection but differences among rootstocks did not always agree with reported field performances. Despite severe biomass reductions plants maintained their relative distribution of biomass among different components of the root system, and no dead roots were observed. Root respiration was reduced by CLas infection and was overall higher in tolerant cultivars suggesting its potential as a physiological marker. This study improves our knowledge about the strengths and weaknesses of assessing rootstock traits of grafted trees in a controlled greenhouse setting. Results from the study suggest that in addition to HLB tolerance, other rootstock traits will ultimately have major contributions to field survival and productivity of the grafted trees in an HLB endemic production environment.
Huanglongbing (HLB) is a major disease of citrus associated with phloem-limited bacteria in the genus Candidatus Liberibacter that affects all known citrus species and relatives, with many commercial cultivars being greatly damaged. Testing cultivar tolerance to HLB in field conditions is difficult because of the erratic spread of the bacteria, scion and rootstock interactions, and influence of many biotic and abiotic factors on the tree response to the disease. This study aimed to validate the effect of CLas infection on different citrus species and hybrids thought to have different levels of tolerance to the disease using CLas graft inoculation under controlled greenhouse conditions. Young potted seedlings from 12 different citrus germplasm selections were graft-inoculated with CLas or mock-inoculated. Plants were monitored periodically during 18 months for canopy growth, HLB and nutritional leaf symptoms, and leaf CLas titers. The leaf nutrient content was measured at the end of the experiment. Roots were also assessed at 18 months after inoculation (mai) for CLas titers and biomass distribution. There were significant differences in most analyzed variables of healthy and infected plants. Some plants of all cultivars were successfully infected; however, overall, the CLas transmission rate was low and inconsistent. Ct values of roots were generally higher than those in leaves at 18 mai. HLB symptoms were not observed on seedlings until 1 year after inoculation; at 18 mai, infected trees of all cultivars were HLB symptomatic. Significant shoot and root biomass reductions (44%–75%) in infected ‘Cleopatra’, ‘Duncan’, ‘Olinda Valencia’, ‘Sunburst’, and ‘Valencia 1-14-19’, considered susceptible to HLB, were measured. These cultivars also showed more severe HLB symptoms than the presumed tolerant cultivars such as Microcitrus inodora, Rich 16-6 trifoliate orange, and US-897. This study provides new knowledge of the efficacy and value of greenhouse screening of citrus germplasm for response to HLB to support the development of new cultivars with improved HLB tolerance or resistance.
The citrus industry in Florida faces a destructive endemic disease, known as huanglongbing (HLB), associated with Candidatus Liberibacter asiaticus (CLas), a phloem-limited bacterium, and transmitted by the Asian citrus psyllid (ACP). Rootstocks are regarded as critical to keep citrus production commercially viable and help trees cope with the disease. Although most scions are susceptible, some rootstocks are HLB-tolerant and may influence ACP infestation and CLas colonization and therefore the grafted tree tolerance. This study aimed to elucidate the relative influence of rootstock and scion on insect vector infestation and CLas colonization under natural HLB-endemic conditions. Seven commercial rootstock cultivars with different genetic backgrounds were grafted with ‘Valencia’ sweet orange (Citrus sinensis) or were self-grafted (non-‘Valencia’) and planted in an open field where ACP and CLas were abundant. ACP infestation was determined weekly during periods of leaf flushing, and leaves and roots were analyzed every 3 months to determine CLas titers. Trees with ‘Valencia’ scion were more attractive to the psyllids than non-‘Valencia’ scions. This was also associated with a higher number of bacteria and a larger abundance of foliar HLB symptoms. The influence of the rootstock on the psyllid attraction of grafted ‘Valencia’ scion was less evident, and leaf CLas titers were similar regardless of the rootstock. Among the non-‘Valencia’ scions, Carrizo had the lowest and US-942 the highest leaf CLas titers. Root CLas titers also varied among cultivars, and standard sour orange roots harbored more bacteria than some trifoliate orange hybrid rootstocks such as US-942. In some trees, CLas was detected first in the roots 4 months after planting, but root CLas titers remained low throughout the study. In contrast, leaf CLas titers increased over time and were considerably higher than root titers from 7 months until the end of the study, 15 months after planting. Overall, the results of this study demonstrate a greater relative influence of the scion than the rootstock on ACP infestation and CLas colonization during the early stages of infection. This suggests that other cultivar-specific traits, such as the ability to tolerate other stresses and to absorb water and nutrients more efficiently, along with influences on the scion phenology, may play a larger role in the rootstock influence on the grafted tree tolerance during the later stages of HLB progression.
Most of the commercially important citrus scion cultivars are susceptible to Huanglongbing (HLB), which is the most devastating disease the citrus industry has ever faced. Because the rootstock can influence the performance of the scion in various ways, including disease and pest tolerance, use of superior rootstocks can assist citrus growers with minimizing the negative effects of HLB. The objective of this study was to assess rootstock effects on the horticultural performance and early production potential of ‘Hamlin’ sweet orange (Citrus sinensis) trees in commercial field settings under HLB-endemic conditions. Two field trials were conducted in different locations in Central and Southeast Florida. The trials were established in 2015 and included 32 diverse diploid and tetraploid rootstock cultivars and advanced selections. One trial was performed in Highlands County, FL, on a poorly drained flatwoods-type site. Another trial was performed in Polk County, FL, on a well-drained sandy Central Florida Ridge site. Horticultural traits including tree height, canopy volume, trunk diameter, canopy health, leaf nutrient content, yield, and fruit quality were assessed during the 2018–19 and 2019–20 production years. Significant differences were found among trees on different rootstocks for most of the measured traits, particularly tree vigor and productivity, but rootstock effects also varied by location. Rootstocks that induced large tree sizes, such as the diploid mandarin × trifoliate orange hybrids ‘X-639’, ‘C-54’, ‘C-57’, and ‘C-146’, also induced higher yield, but with lower yield efficiency. Most of the tetraploid rootstocks significantly reduced tree size, among which ‘Changsha+Benton’, ‘Green-3’, ‘Amb+Czo’, ‘UFR-3’, and ‘UFR-5’ induced high yield efficiency. Therefore, these rootstocks have the potential to be used in high-density plantings. However, trees on some of these small size-inducing rootstocks had a higher mortality rate and were more vulnerable to tropical force winds. This study provides important information for the selection of rootstocks with the greatest production potential in an HLB-endemic environment, especially during the early years of production.