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  • Author or Editor: William R. Graves x
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Development of half-sib Gleditsia triacanthos inermis Willd. (honey locust) seedlings was studied during exposure to osmotic and high root temperature stress. Seven days after seed scarification, seedlings of uniform fresh weight were transferred to static hydroponic culture vessels in a growth chamber. Three days later, vessel solutions were replaced with polyethylene glycol 8000-amended solutions with osmotic potentials (ψπ) of -0.05, -0.10, or -0.20 MPa at 23C. Within each ψπ treatment, root temperature was increased from ambient (23C) to 35C for 0, 6, 12, or 24 hr day-1 for 20 days. Root and shoot dry weights decreased with increasing exposure to 35C among seedlings in the -0.05 MPa solution and decreased for seedlings in -0.10 and -0.20 MPa solutions in all temperature regimes. Epicotyl expansion tended to decrease with decreasing ψπ and increasing exposure to 35C. However, for plants in the -0.20 MPa solution, epicotyl length was greatest when roots were exposed to 35C for 6 hr day-1.

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Alnus maritima (Marsh.) Nutt. (seaside alder) is a rare, woody-plant species with potential for use in managed landscapes. Information on the propagation and production of this species is not available. Our objective was to evaluate the use of softwood cuttings to propagate A. maritima, with emphasis on how indole-3-butyric acid (IBA), plant provenance, and time of collection affect cutting survival, rooting percentage, the number of roots produced, and their length. Propagation trials were conducted with cuttings from seven trees on the Delmarva Peninsula (Eastern Shore of Maryland and southern Delaware) and seven trees in Oklahoma. Cuttings from mature plants in both provenances were collected on 14 June and 23 Aug. 1998; wounded; treated with IBA at 0, 1, or 8 g·kg-1; and placed under intermittent mist in a greenhouse for 9 weeks. Use of IBA at 8 g·kg-1 caused a greater rooting percentage (68%), root count (7.2), and root length (39.2 mm) than did the other IBA rates when applied to cuttings from Oklahoma in June, but IBA had little effect on cuttings from the Delmarva Peninsula. Across IBA treatments, rooting of cuttings from Oklahoma (55% in June and 12% in August) was greater than the rooting of cuttings from Delmarva (27% in June and 3% in August). Cuttings from Oklahoma had greater survival, callus development, root length, and root count than did cuttings from the Delmarva Peninsula during June and August trials. Averaged over IBA treatments and provenances, cuttings collected on 14 June rooted more frequently (41%) than did cuttings collected 23 Aug. (8%). We conclude that softwood cuttings from mature plants are an effective way to multiply clones of A. maritima, particularly when cuttings are collected early in the season and treated with IBA at 8 g·kg-1.

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Horticulturists have not promoted use of Dirca palustris L. (eastern leatherwood) despite its suite of traits valued by gardeners and landscapers. Horticultural production of D. palustris may be hindered by slow shoot growth and sensitivity of plants to edaphic conditions. Because of discrepancies in reported tolerances of D. palustris to root-zone pH, we assessed whether pH of soils supporting indigenous populations in Florida, Maine, and North Dakota corresponded to responses of seedlings from the three provenances to root-zone pH of 4.5 to 7.3 in soilless media. Regression showed that root zones at pH 5.8 promoted maximum stem length of seedlings from Florida and North Dakota, whereas root zones at pH 4.5 led to maximum stem length of seedlings from Maine. Root-zone pH 5.6 and 5.5 fostered maximum root and shoot dry weight, respectively, for seedlings from Florida, whereas root zones at pH 4.5 promoted maximum root and shoot dry weights of seedlings from Maine and North Dakota. Averaged over provenance, relative leaf greenness decreased by 62%, and foliar nitrogen, iron, manganese, and zinc decreased by 49%, 70%, 95%, and 48%, respectively, as root-zone pH increased from 4.5 to 7.3. Foliar phosphorus decreased at both low and high pH. The pH of soils where seeds were collected did not predict optimal root-zone pH for stem length or biomass accrual in soilless media; genotypes from soils with a pH of 7.4 in North Dakota did not exhibit greater tolerance to high pH than genotypes from Maine or Florida, where pH of indigenous soil was 6.1 and 5.2, respectively. Averaged over pH treatments, seedlings from Florida showed the greatest stem length and formed the most shoot biomass, whereas seedlings from North Dakota had stouter stems, greater root biomass, and greater root-to-shoot ratios than did seedlings from Florida and Maine. Our results illustrate that acidic media facilitate horticultural production of D. palustris, that further evaluation of provenance differences could facilitate selection of genotypes for horticulture, and that tolerances of genotypes to root-zone pH do not strictly correspond to the pH of soils on which they were indigenous.

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The genus Nyssa L. includes several woody species with traits valued by horticulturists, but only black gum (Nyssa sylvatica Marsh.) is prevalent in the nursery trade. A congener, swamp tupelo (Nyssa biflora Walt.), might be a marketable shade tree, but little is known about propagating it from seeds. Because cold, moist stratification overcomes embryo dormancies of black gum, we compared germination of cleaned seeds (surrounding pulp of fruits removed) of swamp tupelo and black gum that were stratified at 5 °C for 0, 14, 28, 42, 56, 70, 84, and 112 days. Seeds of swamp tupelo within intact drupes were also stratified. Across all durations of stratification, 79% of cleaned seeds of swamp tupelo germinated, whereas 11% of seeds within drupes germinated. Germination value of cleaned seeds of swamp tupelo increased from 1.26 to 3.23 as duration of stratification increased. Although cleaned seeds of black gum responded similarly, the benefit of stratification was more pronounced, and the mean germination percentage was lower than for swamp tupelo (66% vs. 79%). In a second experiment, irrigation with low and high concentrations of an extract of fruit pulp of swamp tupelo reduced germination of seeds of basil (Ocimum basilicum L. ‘Superbo’), spinach (Spinacea oleracea L. ‘Bloomsdale’), zinnia (Zinnia ×marylandica Spooner, Stimart, and Boyle ‘Double Zahara Cherry’), and swamp tupelo by 25% to 63% (low concentration) and 40% to 70% (high concentration). Propagators should remove the surrounding pulp from seeds of swamp tupelo and cold stratify them at least for 4 weeks.

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Two species of North American shrubs in the genus Ptelea (Rutaceae), commonly known as eastern and western hoptree or wafer ash, have unfulfilled potential to increase the diversity of managed landscapes and support populations of pollinators and swallowtail butterflies. The white flowers of Ptelea are highly fragrant, and pistillate flowers give rise to clusters of distinctive samaras. The insufficiency of information about improving germination of seeds of Ptelea trifoliata and lack of recommendations for Ptelea crenulata prompted us to investigate effects of pericarp removal and cold (4 °C) stratification periods of 0, 4, 8, and 16 weeks on the germination of seeds of these two species. Samaras were collected from multiple plants of both species in the midwestern United States (P. trifoliata) and California (P. crenulata). The germination percentage of viable seeds, calculated after tests of viability with tetrazolium chloride of seeds that did not germinate, increased with longer stratification periods, with 100% germination for P. crenulata and 91% germination for P. trifoliata after 16 weeks of stratification. The germination value, a measure of the speed and uniformity of germination, and peak value also increased with longer stratification in both species. Pericarp removal increased the germination percentage of both species and increased the peak and germination value of P. crenulata. Propagators seeking to grow these species of Ptelea from seed should remove the pericarp and cold-stratify seeds for 16 weeks to improve germination success.

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Numerous genera of plants are distributed in both eastern North America and in portions of California with the dry summers of Mediterranean climates. We compared effects of flooding and drought on relative growth rate (RGR), photosynthesis, and biomass of seedlings of two genera, Sambucus L. and Ptelea L., with congeners in both regions. Ptelea crenulata Greene and Sambucus nigra ssp. cerulea (Raf.) R. Bolli from the San Francisco Bay area and Ptelea trifoliata L. and Sambucus nigra ssp. canadensis (L.) R. Bolli from mesic, deciduous forests in eastern North America were studied. Potted seedlings were subjected to six treatments, three extents of root-zone inundation and three severities of drought (irrigation when soil moisture by volume decreased to 5%, 10%, and 20%). After 5 weeks, deleterious effects of inundation were more pronounced among plants from the West than among their eastern congeners. For example, RGR of western and eastern Sambucus with completely inundated root zones was reduced 116% and 25%, respectively. All western and eastern Ptelea died when root zones were completely inundated, but inundating the lower half of the root zone killed all western plants but only reduced RGR among eastern plants. Photosynthesis of Sambucus from the West was lowest with complete inundation and was similar across the remaining treatments. In contrast, photosynthesis of eastern Sambucus was lowest during severe drought but otherwise similar. Photosynthesis of Ptelea was sensitive to both drought and flooding, and moderate root-zone water content led to the highest rates for both western and eastern plants. For both genera, maximal photosynthesis per unit leaf area was greater among western than eastern plants, but eastern plants had greater total leaf area and biomass. Root-to-shoot ratios of western Sambucus were greater than ratios of plants from the East after all treatments, whereas western Ptelea had greater root-to-shoot ratios than eastern Ptelea only under severe drought. Although comparative sensitivity to drought of plants from California and eastern North America varied in these genera, Mediterranean Sambucus and Ptelea both showed greater sensitivity to root-zone inundation than did their eastern congeners.

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A reputation for coarse root systems with dominant taproots, and for slow shoot development among seedlings, limits use of hickory species (Carya Nutt.) that could increase diversity in managed landscapes. We examined effects of root pruning and application of auxin on root and shoot development of seedlings of several species of hickory. Our hypothesis was that pruning the radicle shortly after seed germination and subsequent treatment with auxin would increase root branching without curtailing development of the shoot. Germinated seeds of Carya aquatica (F. Michx.) Nutt., Carya cordiformis (Wangenh.) K. Koch, Carya laciniosa (F. Michx.) Loudon, Carya ovata (Mill.) K. Koch, and Carya tomentosa (Lam.) Nutt. were treated by removing two-thirds of the length of the radicle with and without immediate application of 3000 mg·L−1 indole-3-butyric acid (IBA) via Hormex rooting powder #3 to the remaining one-third of radicle. Neither treatment altered stem height, stem caliper, or root dry weight. After 75 days, root-pruned seedlings of Carya ovata without auxin had 42% fewer fibrous first-order lateral roots than did unpruned controls. Root pruning plus auxin led to a 79% increase in the number of fibrous first-order lateral roots of C. laciniosa and an ≈50% increase in the shoot dry weight of C. aquatica. Both root pruning and root pruning plus auxin evoked formation of taproot branches for all species. Because species differed in responses of root and shoot systems to root pruning with and without auxin, the practice should be implemented cautiously based on the species.

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Symbiotic associations between Alnus maritima (Marsh.) Muhl. ex Nutt. (seaside alder) and actinomycetes in the genus Frankia Brunchorst result in root nodules in which atmospheric nitrogen (N) is fixed. The economic and environmental benefits of N fixation have led to interest in inducing root nodules during production of A. maritima. Because woody plants produced in nurseries typically are provided N fertilizer, our objectives were to determine how applied N influences nodulation of A. maritima and to characterize how short-term changes in root-zone N affect the function of nodules. Potted seedlings were grown in perlite that was inoculated with 30 mL of soil from the root zones of mature plants in their native habitat on the Delmarva Peninsula. Each pot was drenched once daily for 10 weeks with nutrient solution that contained ammonium nitrate at 10 concentrations from 0 to 8 mm. Plants that received no ammonium nitrate formed the most nodules, and nodulation decreased linearly as ammonium nitrate increased from 0.25 to 4 mm. Plants treated with ammonium nitrate at 4 or 8 mm formed nearly no nodules, while ammonium nitrate at 0.5 mm resulted in vigorous plants with an average nodule count of 70. In a second experiment, a population of nodulated seaside alders was established by irrigating seedlings in inoculated perlite once daily with 0.5-mm ammonium nitrate for 6 weeks. Plants then were provided ammonium nitrate at 0.5, 2, or 4 mm for 2 weeks. Acetylene-reduction assays showed suppressed nodule activity among plants provided 2- and 4-mm ammonium nitrate. Daily irrigation of those plants with N-free solution subsequently led to a rapid depletion of root-zone N and to a concomitant resurgence of nodule activity. These results demonstrate that N fertilization can be managed to promote nodulation of A. maritima and show that decreased nodule activity caused by short-term increases in root-zone N is reversible.

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Although many species of Alnus Miller grow in wet soils, none is as closely associated with low-oxygen, waterlogged soils as Alnus maritima (Marsh.) Muhl. ex Nutt. (seaside alder). An actinorhizal species with promise for use in horticultural landscapes, land reclamation, and sustainable systems, A. maritima associates with Frankia Brunchorst, thereby forming root nodules in which gaseous nitrogen is fixed. Our objective was to determine how root-zone moisture conditions influence the occurrence, location, and anatomy of nodules on A. maritima. Plants of Alnus maritima subsp. maritima Schrader and Graves were established in root zones with compatible Frankia and subjected to four moisture regimens (daily watered/drained, partially flooded, totally flooded, and totally flooded with argon bubbled through the flood water) for 8 weeks. Oxygen content of the root zone, number and location of nodules on root systems, and dry weight and nitrogen content of shoots were determined. Root-zone oxygen content ranged from 17.3 kPa for daily watered/drained plants to 0.9 kPa for argon-treated plants. Across all treatments, 87% of the nodules were within the upper one-third (4 cm) of the root zone. Although shoot dry weights of daily watered/drained and partially flooded plants were not different, daily watered/drained plants had more nitrogen in their leaves (2.53 vs. 2.21 mg·g-1). Nodulation occurred in all treatments, but nodules on totally flooded roots (with or without argon) were limited to a single lobe; in contrast, multilobed nodules were prevalent on partially flooded and daily watered/drained plants. Frankia infection within submerged nodule lobes was limited to one or two layers of cortical cells. Submerged nodules developed large air spaces between cortical cells, and phenolic-containing cells appeared to inhibit Frankia expansion within the nodule. These data suggest that access to root-zone oxygen is critical to the Frankia-A. maritima subsp. maritima symbiosis, and that plants of this subspecies in the drained soils of managed landscapes may benefit more than plants in native wetland habitats from nodulation and nitrogen fixation.

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Alnus maritima (Marsh.) Muhl. ex Nutt. is a large shrub or small tree with potential for use in managed landscapes. Because the three subspecies of A. maritima are indigenous only to areas with mild winter temperatures (USDA hardiness zones 7a and 7b), knowledge of their cold acclimation and cold hardiness is vital if they are to be used where winters are more harsh. Phenology and depth of cold hardiness were assessed by collecting stem samples seven times from 25 Sept. 2000 to 23 Apr. 2001, subjecting the samples to cold temperature ramping, and determining the lowest survival temperature (LST) via the tissue discoloration method. Samples were collected from indigenous plants of the three subspecies and from plants growing in a common garden near Ames, Iowa (USDA zone 5a). Results indicated that some plants from all three subspecies can survive midwinter extremes as low as -80 °C; that plants grown in Ames achieved a greater depth of cold hardiness for most of the winter and were more uniform in cold hardiness than plants growing in warmer native sites; and that the three subspecies did not differ in phenology or depth of cold acclimation. Results of field trials with plots of 150 plants each installed in three northern hardiness zones (USDA zones 5a, 4a, and 3a) supported these conclusions by showing survival of all 450 plants. We resolved differences among subspecies by rating the percentage of stem tissue survival for each plant in the field plots. Subspecies maritima, from the northernmost provenance (the Delmarva Peninsula), showed the least stem death across all three plots (3.9% tissue death), followed by subsp. georgiensis from northwestern Georgia (10% tissue death), and subsp. oklahomensis from southern Oklahoma (12.8% tissue death). Our results suggest that low temperatures should not limit the use of A. maritima in areas as harsh as USDA zone 3a. Selections based on cold hardiness may allow the use of A. maritima in areas with even colder winters.

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