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  • Author or Editor: Harry T. Horner x
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Knowing whether leguminous trees have the potential to nodulate after infection by rhizobial bacteria is important for managing nitrogen (N) applications during tree production and for culture in the landscape. Although 98% of studied species in the Papilionoideae nodulate, the nodulation status of two tree species in this subfamily is uncertain. Cladrastis kentukea (Dum.-Cours.) Rudd (American yellowwood) did not form nodules during inoculation studies in 1939 and 1992. Nodules were observed on mature Sophora japonica L. (Japanese pagodatree) in Japan and Hawaii in the 1940s, but compatible rhizobia reportedly isolated in Japan are no longer held in bacterial collections. Our objective was to verify further that American yellowwood does not nodulate and to confirm reports that Japanese pagodatree does nodulate. Rhizobia that infect many plant hosts, soil samples and rhizobial isolates from other Sophora spp., and soil samples from mature American yellowwood and Japanese pagodatree were used to inoculate 5-day-old seedlings of American yellowwood, Japanese pagodatree, and control species. Soil from indigenous and introduced trees in the continental United States, Hawaii, Japan, and China was used. Inoculated and uninoculated plants were grown for 7 weeks in sterile Leonard jars or clay pots containing perlite and irrigated with sterile, N-free Hoagland's solution. No inoculation treatment elicited nodulation of American yellowwood or Japanese pagodatree. Our results provide additional evidence that American yellowwood lacks that capacity to nodulate and cast further doubt on nodulation of Japanese pagodatree.

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Early nodulin genes, such as ENOD2, may be conserved and could function as molecular markers for nodulation. Many nodulating and nonnodulating legumes must be analyzed before the role of such genes in nodulation can be determined. Japanese pagodatree and American yellowwood are closely related, ornamental woody legumes. Unsubstantiated reports of nodulation in Japanese pagodatree require confirmation, and American yellowwood has not been observed to nodulate. We investigated the presence of putative ENOD2 genes in these species, and we are studying differential and temporal expression. Genomic DNA of Japanese pagodatree and primers, derived from proline-rich pentapeptide repeats of conserved ENOD2 sequences, were used to obtain a 555-bp PCR fragment. This cloned fragment was used as a probe for Southern and Northern hybridizations. Genomes of Japanese pagodatree and American yellowwood contained sequences that are similar to ENOD2 sequences in other legumes. Treatments with either cytokinin or an auxin transport inhibitor may induce expression of the putative ENOD2 genes. New data on the characteristics of nodulin genes in woody legumes will clarify the nature and evolution of nodulation in legumes and may have implications for developing sustainable nursery production protocols.

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A complete picture of legume nodulation has yet to be elucidated. Most studies of the molecular mechanisms responsible for nodule organogenesis have focused on herbaceous legumes. We investigated the presence of a putative ENOD2 gene and studied the temporal and organ-specific production of its transcripts in an ornamental woody legume, Amur maackia. Primers derived from proline-rich pentapeptide repeats of conserved ENOD2 sequences and the genomic DNA of Amur maackia were used to obtain a 543-bp PCR fragment. Southern and Northern blots were probed with this cloned fragment. The Amur maackia genome contained an ENOD2 sequence that is similar to sequences in other species. Expression of the putative ENOD2 gene was detected in roots, 4 days after rhizobial inoculation, but not in leaves or stems. New data on the characteristics of nodulin genes in woody legumes will be beneficial in clarifying the nature and evolution of nodulation in legumes and may have implications for developing sustainable nursery production protocols.

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ENOD2 and other early nodulin genes are conserved among legumes studied to date and might function as markers for the potential of legumes to nodulate. Early nodulin genes have been characterized only among herbaceous legumes. We are interested in understanding the nature of ENOD2 in a nodulating, woody legume. A 561-bp MaENOD2 PCR fragment was used as a probe to screen a cDNA library from nodules ≈1 mm in diameter on roots of Amur maackia, the only temperate and horticulturally desirable leguminous tree species known to nodulate. Five cDNAs were selected for nucleotide sequence analysis. Sequences were determined by using automated dideoxy sequencing and analyzed for identity to other genes with the Genetics Computer Group (GCG) program. The cDNA clones show 68% to 74% identity at the nucleic acid level with ENOD2 genes of Sesbania rostrata Brem. & Oberm., Glycine max (L.) Merrill, and Lupinus luteus L. Southern and northern analyses are being conducted to investigate the possibility of a gene family and to show differential and temporal production of transcripts, respectively. These studies provide new information about nodulins of woody legumes and are being used to facilitate related research on molecular barriers to nodulation in the closely related, non-nodulating tree species Cladrastis kentukea (Dum.-Cours.) Rudd (American yellowwood) and Sophora japonica L. (Japanese pagodatree).

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Evergreen rhododendrons (Rhododendron L.) are important woody landscape plants in many temperate zones. During winters, leaves of these plants frequently are exposed to a combination of cold temperatures, high radiation, and reduced photosynthetic activity, conditions that render them vulnerable to photooxidative damage. In addition, these plants are shallow-rooted and thus susceptible to leaf desiccation when soils are frozen. In this study, the potential adaptive significance of leaf morphology and anatomy in two contrasting Rhododendron species was investigated. R. catawbiense Michx. (native to eastern United States) exhibits thermonasty (leaf drooping and curling at subfreezing temperatures) and is more winter-hardy [leaf freezing tolerance (LT50) of containerized plants ≈–35 °C], whereas R. ponticum L. (native to central Asia) is less hardy (LT50 ≈–16 °C), and nonthermonastic. Thermonasty may function as a light and/or desiccation avoidance strategy in rhododendrons. Microscopic results revealed that R. ponticum has significantly thicker leaf blades but thinner cuticle than R. catawbiense. There is one layer of upper epidermis and three layers of palisade mesophyll in R. catawbiense compared with two distinct layers of upper epidermis and two layers of palisade mesophyll in R. ponticum. We suggest that the additional layer of upper epidermis in R. ponticum and thicker cuticle and extra palisade layer in R. catawbiense represent structural adaptations for reducing light injury in leaves and could serve a photoprotective function in winter when leaf photochemistry is generally sluggish. Results also indicate that although stomatal density of R. ponticum is higher than that of R. catawbiense leaves, the overall opening of stomatal pores per unit leaf area (an integrated value of stomatal density and pore size) is higher by approximately twofold in R. catawbiense, suggesting that R. catawbiense may be more prone to winter desiccation and that thermonasty may be a particularly beneficial trait in this species by serving as a desiccation-avoidance strategy in addition to a photoprotection role.

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High temperatures are reported to promote day-neutral strawberry (Fragaria ×ananassa) vegetative growth and development and inhibit floral and fruit development, thereby imposing geographic and temporal limitations on fruit production. Day-neutral strawberry response to air temperature has been researched, but specific responses to temperature in the root zone have not. In a 1998 greenhouse experiment, 60 `Tristar' plants were grown hydroponically in a system of individual, temperature-controlled pots. A randomized complete-block design with constant root-zone treatments of 11, 17, 23, 29, and 35 °C and 12 replications were used. Stomatal conductance and transpiration rate were significantly lower for plants at 35 °C, compared with plants at all other temperatures. Leaf area and leaf dry mass of plants at 35 °C were five and four times smaller, respectively, than the combined mean for plants in all other treatments. Leaf area of runner tips was 450 and 44.5 cm2 at 11 and 35 °C, respectively, compared with that of plants at all other temperatures, 1552.1 cm2. Fruit dry mass was 14.5, 21.6, 25.5, 29.0, and 3.96 g per plant at 11, 17, 23, 29, and 35 °C, respectively. Root dry mass was highest at 11 and 17 °C and lowest for plants at 35 °C. The number of flowers, fruit, and inflorescences per plant was reduced at 35 °C, as were individual berry fresh mass and diameter. Overall, `Tristar' growth and development were near optimal at 17, 23, and 29 °C.

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Temperature, as a potential environmental stressor, interacts with photoperiod in floral initiation of June-bearing strawberries (Fragaria ×ananassa), such that high-temperature exposure can result in poor floral initiation. Our objectives were to examine the effects of various durations of high root-zone temperature on floral initiation and development and on vegetative growth and development. In a 1998 greenhouse experiment, hydroponically grown `Allstar' June-bearing strawberry plants were subjected day/night temperatures of 31/21 °C in the root zone for one, two, or three continuous periods (of ≈7 days), followed by exposure to 17 °C for the duration of the experiment. Control plants were raised at 17 °C in the root zone throughout the experiment. An additional temperature treatment was exposure to 31/21 °C in the root zone for two periods, each followed by a period at 17 °C. Plants were arranged in a randomized complete-block design with factorial treatments of duration of high root-zone temperature and harvest time. At the end of each period, plants were harvested and the apical meristems dissected for microscopic evaluation of vegetative and floral meristems and the stage of development of the primary flower. We observed floral initiation in all treatments after photoperiodic induction. However, exposure to 31/21 °C in the root zone during key periods of floral initiation in June-bearing strawberry may alter floral development.

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Early nodulin genes, such as ENOD2, play a role in the first stages of nodulation. Although ENOD2 is conserved among nodulating legumes studied to date, its occurrence and activity have not been studied among woody legumes such as Maackia amurensis Rupr. & Maxim. Our objective was to localize MaENOD2 transcripts during nodule development and describe the anatomy of nodules formed on the roots of M. amurensis in relation to ENOD2 mRNA accumulation. Nodules (<1 mm, 1-2 mm, >2 mm in diameter, and mature) were prepared for light microscopy, sectioned, and stained with safranin and fast green for structural contrast or with the periodic acid Schiff's reaction for starch. The location of ENOD2 transcripts was determined by using in situ hybridization with DIG-labeled sense and antisense RNAs transcribed from a 602-bp fragment of the coding region of MaENOD2. Mature nodules from M. amurensis possessed peripheral tissues, a distal meristem, and a central infected region characteristic of indeterminant development. In situ hybridization showed that MaENOD2 transcripts accumulated in the distribution layer and uninfected cells of the central symbiotic region. Amyloplasts that contained starch grains were identified in these tissues and in the inner parenchyma of the nodule. Throughout nodule development, transcripts were restricted to areas with high levels of stored starch that surrounded cells actively fixing N2. Our results suggest that ENOD2 in M. amurensis may be a cell wall component of tissues that regulate nutrient flow to and from sinks, such as symbiotic regions of a nodule. These data may lead to a better understanding of the role of the ENOD2 gene family during nodulation.

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