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  • Author or Editor: William R. Graves x
  • Journal of the American Society for Horticultural Science x
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Information on the heat resistance of silver maple (Acer saccharinum L.) could help develop stress-resistant Freeman maples (Acer ×freemanii E. Murray). Our first objective was to determine how 26, 30, 32, 34, and 36 °C in the root zone affect growth and water relations of plants from rooted cuttings of a silver maple clone indigenous to Mississippi (33.3 °N latitude). Fresh mass increased over time for plants at all temperatures and was highest for plants with root zones at 30 °C. Quadratic regression functions predicted maximal plant dry mass, leaf surface area, and stomatal conductance at 29, 29, and 28 °C, respectively. Stem xylem water potential (ψ) during the photoperiod decreased linearly with increasing root-zone temperature from -0.83 MPa at 26 °C to -1.05 MPa at 36 °C. Our second objective was to compare six clones of silver maple from the Mississippi location with six clones from 44.4 °N latitude in Minnesota for effects of 35 °C in the root zone on plant growth, stomatal conductance, and stem ψ. Provenance and temperature main effects were significant for most dependent variables, but there were no provenance × temperature interactions. Over both provenances, plant fresh and dry mass, leaf surface area, stomatal conductance, and stem ψ during the photoperiod were higher at 29 than 35 °C. Over both temperatures, plants from Minnesota clones had higher fresh and dry mass and more leaf surface area than plants from Mississippi clones. The lack of temperature × provenance interactions suggests that ecotypic or clinal variation in heat resistance is minimal and will not be useful for identifying superior genotypes for use in interspecific crosses with red maple (Acer rubrum L.).

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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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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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Selection of sugar maples (Acer saccharum Marsh.) and black maples (Acer saccharum Marsh. ssp. nigrum Desm. or Acer nigrum Michx. f.) that will be more resilient than existing cultivars in managed landscapes could be facilitated by defining relationships between geographic origin and foliar traits critical to leaf function. We examined variation in leaf morphology and anatomy of both taxa, known collectively as hard maples, near 43 °N latitude and tested for relationships between foliar traits and the longitude of origin from 70 ° to 94 °W longitude. Leaves exposed to direct solar radiation were sampled from up to 20 trees indigenous at each of 42 sites during 1995 and 1996. All leaves from east of 75.84 °W and from 92.73 °W and further west expressed morphological characters associated with sugar maple and black maple, respectively; leaves with intermediate traits were found between these two longitudes. Leaves from 90 ° to 94 °W had the highest surface area due to increases in the areas of middle and proximal portions of laminae. Up to 1162 trichomes/cm2 were present on the abaxial surface of laminae from west of 85 °W, while laminae from further east were glabrous or had ≤300 trichomes/cm2. Laminae from western habitats also had relatively high stomatal frequency, and stomatal apertures of laminae west of 91 °W were particularly narrow. Longitude did not affect specific weight and thickness of laminae, which averaged 5.5 mg·cm-2 and 90 μm, respectively. Principal component analysis of laminar traits showed existence of two clusters. A large group dominated by data from trees in New England also contained data from trees as far west as ≈93 °W longitude; data for trees further west were clustered separately. Although phenotypic continua were defined, laminae west of 93 °W were distinct, which suggests trees selected there may function differently in managed landscapes than trees selected from native populations further east.

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Responses of five bottomland tree taxa to drought and flooding were studied to identify those adapted to urban environments. During one experiment, containerized `Franksred' red maple [Acer rubrum L. `Franksred' (trademark = Red Sunset)], sweetbay magnolia (Magnolia virginiana L.), black tupelo (Nyssa sylvatica Marsh.), bald cypress [Taxodium distichum (L.) Rich.], and pawpaw [Asimina triloba (L.) Dunal.] were treated with various irrigation regimes for up to 118 days. Net assimilation rate (NAR) and relative growth rate (RGR) were reduced more by flooding than by drought for plants of all taxa, except pawpaw, which showed similar NAR and RGR during flooding and drought. Only sweetbay magnolia and bald cypress maintained positive NAR and RGR during flooding, and sweetbay magnolia was the only taxon that did not produce significantly less leaf surface area, shoot dry mass, and root dry mass during flooding and drought. Apparent morphological mechanisms of stress resistance included an increase in specific mass of leaves (mg·cm-2) during drought for red maple and bald cypress and a 385% increase in the root: shoot mass ratio for droughted plants of pawpaw. Leaf water relations of drought- and flood-stressed `Franksred' red maple and sweetbay magnolia were determined in a second experiment. Predawn and mid-day leaf water potential (ψ) decreased with decreasing root-zone matric potential for both taxa, and transpiration rate was reduced by drought and flooding. Pressure-volume analysis showed that leaves of `Franksred' red maple responded to drought by shifting symplastic water to the apoplast. Leaves of drought-stressed sweetbay magnolia adjusted osmotically by reducing osmotic potential (ψπ) at full turgor by 0.26 MPa. Our results suggest that sweetbay magnolia and bald cypress will perform well at urban planting sites where episodes of drought and flooding regularly occur.

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Alnus maritima (Marsh.) Muhl. ex Nutt. is a rare woody plant species that exists as three subspecies found in widely disjunct locations in the United States. Although there is a growing interest in the phytogeography, ecology, conservation, and landscape potential of this species, the phylogeny of A. maritima has not yet been resolved by using molecular methods. We have combined a relatively new method of genome fingerprinting, ISSR-PCR, and the automated imaging capabilities of GeneScan technology to investigate the molecular systematics of A. maritima. Based on the molecular evidence from 108 ISSR loci, we confirm that the three disjunct populations of A. maritima have diverged sufficiently to be classified as subspecies. Our molecular phylogeny of the three subspecies of A. maritima agreed in topology with a phylogeny produced from morphological data and showed that subsp. oklahomensis is the most distinct of the three subspecies and was the first to diverge. The simultaneous analysis of molecular and morphological data provides a detailed and balanced phylogeny reconstruction for the three subspecies. Our results support the theory that A. maritima originated in Asia, migrated into North America across the Bering land bridge, and was established over a large range in the New World before being forced into its present meager disjunct distribution.

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Genotypic variation and horticultural potential of Alnus maritima [Marsh.] Nutt. (seaside alder), a large shrub or small tree found naturally in only three small, disjunct populations, have not been studied. We examined effects of population of origin and environment on seed germination and growth and morphology of seedlings. The first experiment showed that 6 weeks of cold stratification optimized germination of half-sibling seeds from Oklahoma at 73.2%. When this treatment was applied to multiple groups of half-siblings from all populations in a second experiment, seeds from Oklahoma had a higher germination percentage (55.0%) than seeds from Georgia (31.4%) and the Delmarva Peninsula (14.7%). In a third experiment, morphology and growth of multiple groups of half-siblings from all three populations were compared in one environment. Leaves of seedlings from Oklahoma were longer (12.8 cm) and more narrow (2.15 length to width ratio) than leaves of seedlings from Georgia (12.0 cm long; ratio = 1.76) and the Delmarva Peninsula (11.6 cm long; ratio = 1.86). Seedlings from Oklahoma and Georgia accumulated dry weight at higher rates (181 and 160 mg·d-1, respectively) than seedlings from Delmarva (130 mg·d-1), while seedlings from Oklahoma and Delmarva were more densely foliated (0.72 and 0.64 leaves and lateral shoots per centimeter of primary stem, respectively) than those from Georgia (0.46 per cm). These differences indicate genetic divergence among the three disjunct populations and the potential to exploit genetic variation to select horticulturally superior A. maritima for use in managed landscapes.

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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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Growth of tomato (Lycopersicon esculentum Mill.) plants decreases at root-zone temperatures (RZTs) >30 °C, but no research has been conducted on the effects of changes in root respiration on P acquisition at supraoptimal RZT. We monitored the changes every 3 to 5 days in root respiration, root surface phosphatase activity, and P acquisition of `Jet Star' tomato plants grown in Hoagland's no. 1 solution held at 25 and 36 °C RZT for 19 days. Root respiration rate in plants grown at 25 °C increased linearly from RZT initiation to day 12, but there was no difference in respiration between days 12 and 19. Root respiration at 36 °C, however, increased from RZT initiation to day 8 and then decreased. Shoot P concentration and root phosphatase activity for plants grown at 25 °C did not change during the experiment. Shoot P concentration for plants at 36 °C, however, linearly decreased over time, and root phosphatase activity linearly increased over time. Decreased shoot growth and demand for P along with decreased root respiration after day 8 probably resulted in the decreased P uptake and shoot P concentration in plants grown at 36 °C RZT.

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Carolina buckthorn [Rhamnus caroliniana Walt. or Frangula caroliniana (Walt.) Gray] is an attractive and water-stress-resistant shrub or small tree distributed extensively in the southeastern United States that merits use in managed landscapes. Due to substantial climatic differences within its distribution (30-year normal midwinter minima range from 13 to -8 °C), selection among provenances based on differences in cold hardiness is warranted. Before selections are marketed, the potential of carolina buckthorn to be invasive also merits investigation. Ecological problems resulting from the introduction of Rhamnus L. species in the United States, most notably the dominance of R. cathartica L. (common buckthorn) over neighboring taxa, are due in part to early budbreak. Consequently, we investigated depth of cold hardiness and vernal budbreak of carolina buckthorn and common buckthorn. Stem samples of carolina buckthorn and common buckthorn collected in midwinter survived temperatures as low as -21 and -24 °C, respectively. Although the cold hardiness of carolina buckthorns from Missouri was greater than that of carolina buckthorns from Ohio and Texas on 2 Apr. 2003, there were no differences in cold hardiness of stems from Missouri and Texas on all three assessment dates in the second experiment. All plants survived at both field locations except for the carolina buckthorns from southern Texas planted in Iowa, which showed 0% and 17% survival in 2003 and 2004, respectively. Budbreak of both species with and without mulch in Ames, Iowa, was recorded from 9 Apr. to 10 May 2002. Mean budbreak of common buckthorn was 5.7 days earlier than budbreak of carolina buckthorn, and buds of mulched carolina buckthorns broke 4.2 days earlier than did buds of unmulched carolina buckthorns. We conclude that the cold hardiness of carolina buckthorn is sufficient to permit the species to be planted outside of its natural distribution. Populations of carolina buckthorn in Ohio and Missouri should be the focus of efforts to select genotypes for use in regions with harsh winters. Phenology of its budbreak suggests carolina buckthorn will not be as invasive as common buckthorn, but evaluation of additional determinants of invasiveness is warranted.

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