WINTER COLD HARDINESS OF SEVEN WILDLAND SHRUBS R. H. Walser D.J. Weber E. D. McArthur S. C. Sanderson ABSTRACT supercooling of water (ice formation avoidance) is involved (George and others 1974). Death occurs when ice crystals form within the cells and rupture cellular membranes in this manner; ice, not low temperatures, causes death. Plants can also deharden following warm spells during the winter period. Howell and Weiser (1970) found that short-term changes in cold resistance of apple stems were closely related to the air temperatures of the preceding day. Hardy plants dehardened during the winter as much as 15 oc in 1 day in a warm greenhouse, and rehardened 15 oc in 3 days when they were held at -15 °C. The maximum cold hardiness achieved by 'Haralson' apple bark was approximately -50 °C in midwinter and -7 °C in the early fall and at bloom time in the spring. Thus, a cold spell in early fall or spring or severe cold in midwinter following a warm spell could cause considerable damage. Various methods have been used in the laboratory to determine shoot and bud cold hardiness. Stergios and Howell (1973) evaluated several viability tests for coldstressed plants. They found that the tissue browning test was the most reliable, but required considerable time and was qualitative. In this test, 15-cm-long stem sections were placed in a chamber where the temperature was lowered 10 °Cih. Samples were removed at 5 oc intervals and placed in a humid chamber at room temperature for 5 days to incubate, they then were examined for damage. Undamaged bark remained green, while damaged bark first became water soaked, then darkened, and ultimately became soft. We used this technique to evaluate seven wildland shrubs in relation to the temperatures that first cause damage and temperatures that would most likely kill the plant. Winter kill of wildland shrubs can be caused by anumber of factors including freezing. Freezing damage and lethal temperatures were determined and are as follows for rubber rabbit brush (Chrysothamnus nauseosus) (-30 °C), Stansbury cliffrose (Cowania stansburiana) (-55 °C), antelope bitterbrush (Purshia tridentata) (-50 °C), big sagebrush (Artemisia tridentata) (-80 °C), fourwing saltbush (Atriplex canescens) (-80 °C), shadscale (Atriplex confertifolia) (-80 °C) and winterfat (Ceratoides lanata) (-80 °C). In April as the buds began to swell, shoots could be killed at temperatures around -5 °C for rubber rabbitbrush, -15 °C for antelope bitterbrush and big sagebrush, -25 °C for shadscale and Stansbury cliffrose, and -35 for fourwing saltbush and winterfat. oc INTRODUCTION Winter kill of perennial agriculture crops is well documented. On the other hand, wildland shrubs are assumed to be very hardy and few studies have been done to determine their cold hardiness (VanEpps 1975; Jensen and Urness 1979; Hanson and others 1982; Nelson and Tiernan 1983). Wildland shrubs represent a major forage source for livestock and wildlife during the harsh winter period (McArthur 1988). Reports oflarge areas of die-off have increased the interest in the possibility of winter damage as a factor in shrub die-off. Temperate zone woody plants have the ability to acclimate during the fall season and thus withstand extremely cold winter temperatures. Research has shown that cold acclimation is a 2-stage sequence. Photoperiod is the initial stimulus that triggers the various metabolic events leading to cold acclimation, and cold temperature triggers the second state of the process (Howell and Weiser 1970). The mechanisms utilized by plants to avoid freezing injury are not co~pletely understood, but it is generally accepted that MATERIALS AND METHODS One-year-old shoots from rubber rabbitbrush, Stansbury cliffrose, antelope bitterbrush, big sagebrush, fourwing saltbush, shadscale, and winterfat were collected monthly from natural populations in or near Skull Valley, UT, for two winters (1986 and 1987). The shoots were cut into 15-cm-long sections and three sections from each species were bundled together. Eight large bundles were then formed with one bundle from each species and treated as follows: seven bundles were placed in a freezing chamber programmed to lower the temperature 10 °Cih. The samples were removed from the chamber when the predetermined temperatures were reached. The samples were then placed under a mist system in a greenhouse (20 °C) and evaluated for damage (browning of phloem, Paper presented at the Symposium on Cheatgrass Invasion, Shrub DieOff and Other Aspects of Shrub Biology and Management, Las Vegas, NV, April 5-7, 1989. R. H. Walser is Associate Professor, Department of Agronomy and Horticulture and D. J. Weber is Professor, Department of Botany and Range Science, Brigham Young University, Provo, UT 84602; E. D. McArthur is Project Leader and Chief Research Geneticist and S. C. Sanderson is Research Geneticist, Intermountain Research Station, Forest Service, U.S. Department of Agriculture, Shrub Sciences Laboratory, Provo, UT 84606. 115 This file was created by scanning the printed publication. Errors identified by the software have been corrected; however, some errors may remain. buds, or xylem) following 1, 4, and 10 days in the greenhouse. The eighth bundle (control) was placed directly in the greenhouse and evaluated with the treated samples. Shoots were rated for the temperature when the first visible tissue damage was evident and when no visible live tissue was evident and no bud growth occurred following 10 days in the greenhouse. In January of each winter, a double amount of shoots was harvested from the field. One set of shoots was processed as previously described, while the second set of shoots was placed in the greenhouse (20 °C) for 6 days, after which they were exposed to low temperatures as previously described. The purpose of this experiment was to determine the amount of dehardening each of the species was capable of achieving during midwinter. RESULTS AND DISCUSSION The results illustrate the great amount of cold tolerance that is developed by the tested shrubs. In all cases, the lowest temperature to kill the shrub shoots occurred in the coldest part of the winter (January-February) as shown in figures 1 through 4. However, there was a very wide temperature range between the point where first visible damage occurred and where no visible live tissue was evident (for example, -27 oc and -80 oc in big sagebrush). Both temperatures were considerably higher in December and March. Based on our experience with fruit trees and ornamental shrubs, and our observations of the progression of damage and regrowth of the treated shoots, we suggest that the temperature which would cause 0 0 [] • Damaged Killed Rubber Rabbitbrush Stansbury Cliffrose -10 6 !!_.. -20 0 -20 !!_.. Q) ~ ~-40 Cll ia. .a e e -30 Q) E Q) Q) 1- 1- -60 -40 •• --80 -50 Dec Jan Feb Dec Mar Jan Feb Mar Time (months) Time (months) 0 Damaged Killed 0 Big Sagebrush Antelope Bitterbrush -20 -20 E -40 0 !!_.. ~ .a Q) ~ -40 ('(I ; ~a. a. -60 E Q) E 1- Q) 1- -60 -80 • •• -100 Dec Jan Feb Damaged Killed -80 Mar Dec Time (months) Jan Feb Time (months) Figure 1-Winter shoot damage (dark square) and winter plant death (dark triangle) in rubber rabbitbrush (top graph) and big sagebrush (bottom graph). Figure 2-Winter shoot damage (dark square) and winter plant death (dark triangle) in Stansbury cliffrose (top graph) and antelope bitterbrush (bottom graph). 116 Mar -20 -30 •• -20 Damaged Killed Damaged Killed Fourwing Saltbush -30 Wlnterfat -40 -40 0 0 t.. -50 t.. !:::J ~ :::J ; -50 ; i_ Q; 0.. -SO E ~ E ~ -SO -70 -70 • -80 • -80 Dec Jan Feb -90 Dec Mar Feb Jan Mar Time (months) Time (months) Figure 4-Winter shoot damage (dark square) and winter plant death (dark triangle) in winterfat. -20 Damaged Killed Shad scale -30 Winter! at -40 E Shadscale Cll .aas -50 Sattbush Q; 0.. l! E ~ -SO 1:: as a: Sagebrush Cliff rose -70 Bitterbrush -80 Dec Jan Feb Rabbitbrush Mar Time (months) l?LI • -100 Six days warming No warming -80 -60 -40 -20 0 20 Temperature (°C) Figure 3-Winter shoot damage (dark square) and winter plant death (dark triangle) in fourwing saltbush (top graph) and shadscale (bottom graph). Figure 5-Effect of January warming on the cold tolerance of winterfat, shadscale, fourwing saltbush, big sagebrush, Stansbury cliffrose, antelope bitterbrush, and rubber rabbitbrush. enough damage to the plants so that they would not survive the stresses of the following summer would fall close to the midpoint between the two temperatures ranges. That midpoint temperature would be approximately -35 °C in December and -20 oc in March for rubber rabbitbrush, -50 oc and -40 oc for big sagebrush, -47 oc and -36 oc for Stansbury cliffrose, -45 oc and -35 oc for antelope bitterbrush, -55 oc and -47 oc for fourwing saltbush, -55 oc and -47 oc for shadscale, and -57 °C and -45 oc for winterfat. All of the plants tested showed considerable de hardening following the January warm treatment. The first damage and final killing damage temperatures were reduced by 50 percent in almost all of the shrubs (fig. 5). The warming period had the least effect on rubber rabbitbrush. Temperatures in the area where the shoots were collected occasionally reach lows of -25 °C to -30 oc (Stevens and others 1983). The coldest temperature ever recorded in Utah was -45 oc (Stevens and others 1983). Under these extreme temperatures, it appears that considerable damage could occur to the shrubs investigated, particularly if the cold spell was preceded by a few days of temperatures considerably above freezing. Another fact that needs to be considered is that roots and crown area of the plants are less cold tolerant than the shoots (Howell and Weiser 1970). Research performed in Minnesota 117 (Wildung and others 1973) indicated that apple stems could tolerate temperatures of -40 °C, while roots were killed at -14 °C. Thus, a cold spell occurring without a protective snow cover would very likely cause much more damage than with a snow cover. Jensen, C. H.; Urness, P. J. 1979. Winter cold damage to bitterbrush related to spring sheep grazing. Journal of Range Management. 35: 142-145. McArthur, E. D. 1988. New plant development in range management. In: Tueller, P. T., ed. Vegetation science applications for rangeland analysis and management. The Netherlands: Dordrecht: 81-112. Nelson, D. L.; Tieman, C. F. 1983. Winter injury of sagebrush and other wildland shrubs in the western United States. Res. Pap. INT-314. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 17 p. Stergios, B. C.; Howell, C. S., Jr. 1973. Evaluation of viability tests for cold stressed plants. Journal of American Society of Horticultural Science. 98: 325-330. Stevens, D. J.; Brough, R. C.; Griffin, R. D.; Richardson, E. A. 1983. Utah weather guide. West Jordan, UT: Center for Applied Climateology. 46 p. VanEpps, G. A. 1975. Winter injury to fourwing saltbush. Journal of Range Management. 28: 157-159. Wildung, D. K.; Weiser, C. J.; Pellett, H. M. 1973. Temperature and moisture effects on hardening of apple roots. HortScience. 8: 53-55. ACKNOWLEDGMENT This research was supported in part by National Science Foundation Grant PCM-8320462 and was facilitated by a cooperative agreement between Intermountain Research Station (USDA Forest Service) and Utah Division of Wildlife Resources Project W-82-R. REFERENCES George, M. F.; Burke, M. J.; Weiser, C. J. 1974. Supercooling in overwintering azalea flower buds. Plant Physiology. 54: 29-35. Hanson, C. L.; Johnson, C. W.; Wight, J. R. 1982. Foliage mortality of mountain big sagebrush (Artemisia tridentata ssp. vaseyana) in southwestern Idaho during the winter of 1976-77. Journal of Range Management. 35: 142-145. Howell, G. S.; Weiser, C. J. 1970. Fluctuations in the cold resistance of apple twigs during spring de hardening. Journal of American Society of Horticultural Sciences. 95: 190-192. 118