It is not known when changes in primary direct heat stress tolerance of conifer seedlings occur in relation to other seasonally changing physiological parameters. This information should be incorporated into nursery practices and the matching of genotypes to landscape sites. Greenhouse-cultured, container-grown Douglas-fir, Engelmann spruce, and ponderosa pine. were cold acclimated and reacclimated in growth chambers over 19 weeks. Direct heat stress tolerance of needles, cold hardiness, and bud dormancy were measured weekly. Douglas-fir and Engelmann spruce heat stress tolerance increased with the development of new growth through one complete growth cycle, i.e., bud break, maturation, cold hardening, dehardening, and bud break the following growing season. Ponderosa pine differed in that new needles had intermediate tolerance, and fully cold hardy needles were the most intolerant. In none of the species did the timing of changes in heat stress tolerance coincide consistently with changes in cold hardiness or bud dormancy.
Karen E. Burr, Stephen J. Wanner, and Richard W. Tinus
Rajeev Arora, Dharmalingam S. Pitchay, and Bradford C. Bearce
This study evaluated the effect of reversible water stress on heat stress tolerance (HST) in greenhouse-grown geraniums. Water stress was imposed by withholding irrigation until pots reached ≈30% (by weight) of well-watered (control) plant pots, and maintaining this weight for 7 days. Control plants were watered to just below field capacity, every other day. Leaf xylem water potential (LXWP, MPa), leaf-relative water content (LRWC,%), media water content (MWC, % fresh weight), and heat stress tolerance (HST, LT50) were determined for control and stressed plants. HST (LT50), defined as temperature causing half-maximal percent injury, was based on electrolyte leakage from leaf disks subjected to 25 to 60C. Control-watering was restored in stressed plants and above measurements made after 7 days of recovery. Data indicate: 1) LXWP, LRWC, and MWC in control and stressed plants were –0.378 and –0.804 MPa, 92.31% and 78.69% and 82.86% and 15.5%, respectively; 2) HST increased significantly in stressed as compared to control plants (LT50 of 55C vs. 51C); 3) control plants were near maximally injured by 53C treatment and sustained more than 3-fold greater injury than stressed plants at 53C. In recovered plants, LXWP and RWC reversed back to control levels, paralleled by loss of higher HST.
S.M. Hum-Musser, T.E. Morelock, J.B. Murphy, and R.L. Henry
Seed germination of spinach (Spinacia oleracea L.) is partially inhibited by a high germination temperature (35 °C). Tolerance of high germination temperatures varies widely depending on the variety used. We ascertained that seed germination of these spinach varieties was thermoinhibited at 35 °C and secondary dormancy was not induced as seeds germinated when transferred to optimum germination conditions (20 °C). Treatment with 99% oxygen and 10 ppm kinetin significantly increased germination of thermoinhibited varieties at 35 °C. During heat stress, all organisms produce heat shock proteins (HSPs), which may function as molecular chaperons, are possibly required for the development of thermotolerance, and may be crucial for cell survival during heat stress. Western blotting of SDS-PAGE gels using antibodies to various heat shock proteins indicated that spinach varieties with the highest degree of thermotolerance have higher levels of HSP expression than varieties with the lowest degree of thermotolerance during germination. These results suggest that thermotolerance could be further improved, either through a breeding program or possibly by genetic engineering.
Aref A. Abdul-Baki
Nine heat-tolerant tomato [Lycopersicon esculentum (Mill.)] breeding lines, four heat-tolerant cultivars, and four heat-sensitive cultivars were evaluated in the greenhouse under high temperature (39C day/28C night) and in the field. Criteria for heat tolerance included flowering, fruit set, yield, fruit quality, and seed production. Under high-temperature conditions, the group of heat-tolerant lines, the heat-tolerant cultivars, and the heat-sensitive cultivars produced, respectively, the following per plant: flowers, 186, 94, and 55; fruit set 70%, 52%, and 30%; yield, 410, 173, and 11 g; and normal mature fruit, 72%, 37%, and 7%. Yields of heat-tolerant lines under high temperature in the greenhouse ranged from 118% to 31% of their respective yields in the field. Yields of heat-tolerant cultivars were 62% of those in the field. In contrast, yields of heat-sensitive cultivars under high temperature were < 1% of their respective yields in the field. High temperature induced flower abscission, reduced fruit set and yield, and increased the incidence of abnormalities. Major fruit abnormalities with high temperatures included cracks, blossomed rot, watery tissue, and small, immature fruits. Production of viable seeds under the high-temperature regime was severely reduced or totally inhibited regardless of the heat-tolerance level exhibited by the line or cultivar. The failure of heat-sensitive and most heat-tolerant cultivars or lines to produce viable seeds under such a high temperature suggests that a lower level of heat stress than that applied in these experiments could allow the production of enough seeds to test the relationship between heat tolerance in a genotype and its ability to produce viable seeds under high temperature. The results indicate that certain lines have high tolerance to heat and, therefore, could provide valuable sources of plant material for physiological studies to establish the physiological and molecular bases of heat tolerance. Some of the heat-tolerant lines might also serve as excellent germplasm sources in breeding heat-tolerant tomato cultivars.
Xiaozhong Liu and Bingru Huang
Understanding physiological factors that may confer heat tolerance would facilitate breeding for improvement of summer turf quality. The objective of this study was to investigate whether carbohydrate availability contributes to changes in turf quality and root mortality during heat stress in two creeping bentgrass [Agrostis stolonifera L. var. palustris (Huds.) Farw. (syn. A. palustris Huds.)] cultivars, `L-93' and `Penncross', that contrast in heat tolerance. Grasses were grown at 14-hour days and 11-hour nights of 22/16 °C (control) and 35/25 °C (heat stress) for 56 days in growth chambers. Turf quality decreased while root mortality increased under heat-stress conditions for both cultivars, but to a greater extent for `Penncross' than `L-93'. The concentrations of total nonstructural carbohydrate (TNC), fructans, starch, glucose, and sucrose in shoots (leaves and stems) and roots decreased at 35/25 °C. The reduction in carbohydrate concentrations of shoots was more pronounced than that of roots. Shoot glucose and sucrose concentrations were more sensitive to heat stress than other carbohydrates. `L-93' maintained significantly higher carbohydrate concentrations, especially glucose and sucrose, than `Penncross' at 35/25 °C. Results suggest that high carbohydrate availability, particularly glucose and sucrose, during heat stress was an important physiological trait associated with heat-stress tolerance in creeping bentgrass.
Yali He, Xiaozhong Liu, and Bingru Huang
The acclimation of plants to moderately high temperature plays an important role in inducing plant tolerance to subsequent lethal high temperatures. This study was performed to investigate the effects of heat acclimation and sudden heat stress on protein synthesis and degradation in creeping bentgrass (Agrostis palustris Huds.). Plants of the cultivar Penncross were subjected to two temperature regimes in growth chambers: 1) heat acclimation—plants were exposed to a gradual increase in temperatures from 20 to 25, 30, and 35 °C for 7 days at each temperature level before being exposed to 40 °C for 28 days; and 2) sudden heat stress (nonacclimation)—plants were directly exposed to 40 °C for 28 days from 20 °C without acclimation through the gradual increase in temperatures. Heat acclimation increased plant tolerance to subsequent heat stress, as demonstrated by lower electrolyte leakage (relative EL) in leaves of heat-acclimated plants compared to nonacclimated plants at 40 °C. Heat acclimation induced expression of some heat shock proteins (HSPs), 57 and 54 kDa, detected in a salt-soluble form (cystoplasmic proteins), which were not present in unacclimated plants under heat stress. However, HSPs of 23, 36, and 66 kDa were induced by both sudden and gradual exposure to heat stress. In general, total protein content decreased under both heat acclimation and sudden heat stress. Cystoplasmic proteins was more sensitive to increasing temperatures, with a significant decline initiated at 25 °C, while sodium dodecyl sulphate (SDS)-soluble (membrane) protein content did not decrease significantly until temperature was elevated to 30 °C. The results demonstrated that both a gradual increase in temperature and sudden heat stress caused protein degradation and also induced expression of newly synthesized HSPs. Our results suggested that the induction of new HSPs during heat acclimation might be associated with the enhanced thermotolerance of creeping bentgrass, although direct correlation of these two factors is yet to be determined. This study also indicated that protein degradation could be associated with heat injury during either gradual increases in temperature or sudden heat stress.
Rib discoloration in crisphead lettuce (Lactuca sativa) has been successfully induced by applying heat stress. Two studies were conducted to determine the effect of short periods (3 and 5 days) of high temperatures (35/25 °C and 35/15 °C day/night temperatures) at various developmental stages (at heading, and at 1, 2, and 3 weeks after heading) on rib discoloration incidence and severity. Lettuce (cv. Ithaca) was most sensitive to heat stress 2 weeks after heading: applying 35/25 °C or 35/15 °C day/night temperatures for 3 or 5 days resulted on average in 46% of mature heads with rib discoloration symptoms. Stressing plants at earlier or later stages resulted in significantly lower incidences of the disorder, with only 4% to 17% plants showing symptoms. More leaves were affected by the disorder when heat stress was applied 2 weeks after heading than when the stress was applied earlier or later. Night temperature and stress duration had no effect on the incidence and severity of rib discoloration. Up to eight leaves, located between the first and fifteenth leaves acropetal to the cap leaf, showed symptoms. This report establishes a direct relationship between rib discoloration and heat stress, proposes a new method to help lettuce breeders screen germplasm for rib discoloration tolerance, and supports the development of tools for predicting the occurrence of rib discoloration in the field according to meteorological data.
Ahmed A. Tawfik and Jiwan P. Palta
The optimum temperature regime for Solanum tuberosum cv. Russet Burbank is usually 20/15°C day/night. We studied the impact of heat stress (30/25°C, day/night) on the growth of this heat sensitive cultivar under controlled conditions (UW-Biotron). Plants were grown in sandy-loam soil which tested at 1500 Kg/ha Ca. Plants were at the maximum temperature for 6h during the middle of the day with a photoperiod of 14 hrs. All pots received identical amounts of total N (rate: 225 Kg N ha1.). The treatments were: (1) NSN: non-split N (N application 1/2 emergence, 1/2 two wks later): (2) SPN: split-N (1/2 emergence 1/6 at 2, 5 and 8 wks later); (3) SPN+Ca: Split-N+Ca (Ca at 2, 5 and 8 wks after emergence, total Ca from CaNO3 was 113 Kg ha1). Total leaf FWT and DWT was significantly reduced in NS treatment by heat stress at 13 wks as compared to optimum conditions. However, this was not reduced in SPN and SPN+Ca. Under heat stress: (a) SPN + Ca gave the highest leaf FWT and DWT, stomatal conductance, transpiration rate, and leaflet tissue Ca content; (b) Young expanding leaflets gave higher growth rate with SPN and SPN + Ca than NSN; (c) Ca content of mature leaflet decreased progressively in both NSN and SPN but not in SPN + Ca. Our results show that application of Ca and N during heat stress can mitigate stress effects and that maintenance of a certain level of calcium in leaf tissue is important under heat stress.
Cuttings of Dendranthema ×grandiflorum `Paragon' were used as a model system to assess the effects of root heating on disease severity. Roots were exposed to single episodes of heat stress, after which they were inoculated with zoospores of Phytophthora cryptogea Pethyb. & Laff. Root damage resulting from heat stress, or heat stress plus Phytophthora, was quantified 5 to 7 days after treatment. Roots of hydroponically grown plants, immersed for 30 min in aerated, temperature-controlled nutrient solutions, were severely damaged at 45C or above. Relatively little phytophthora root rot developed on inoculated plants exposed to 25 or 35C, but infection was severe in roots heated to 40C. Plants grown in potting mix were exposed to heat stress by plastic-wrapping the containers in which they were growing and placing them in heated water baths until roots achieved desired temperatures for 30 min. This system heated roots more slowly than in the hydroponic experiments, and 45 and 50C were less damaging. The amount of Phytophthora-induced root damage was insignificant in containerized plants heated to 25 or 35C, but was highly significant in those heated to 40C or higher. In field experiments, plants were positioned so their containers were either fulIy exposed to the late afternoon sun or heavily shaded to prevent sun exposure. The root zones of sun-exposed pots heated to 45 to 47C, while those of shaded pots never exceeded 34 to 36C. There was a large, highly significant increase in phytophthora root rot severity in the sun-exposed pots compared to shaded plants. These experiments showed that temperatures of 40C or higher, which commonly occur in container-grown plants exposed to solar radiation, can predispose chrysanthemum roots to severe Phytophthora infection.
D.H. Willits and M.M. Peet
Chlorophyll fluorescence was measured under both laboratory and greenhouse conditions in an effort to develop a quick, reliable, and inexpensive laboratory procedure capable of predicting heat stress experienced by tomato (Lycopersicon esculentum Mill.) under greenhouse conditions. The laboratory tests consisted of measurements of the ratio of variable to maximal chlorophyll fluorescence (Fv/Fm) performed on leaf discs taken from whole tomato leaves and placed on a temperature controlled plate. Comparisons were made with greenhouse measurements of the same parameter conducted on intact leaves of whole plants exposed to different temperature treatments imposed by manipulation of the aerial environment of the greenhouse. Dark adaption periods ranging from 15 min to all day in the greenhouse and temperature exposure periods ranging from 5 min to 60 min in the laboratory were compared to find the best correlation between the two tests. Best agreement was obtained with 60 min treatment times in the laboratory and 60 min dark adaption periods in the greenhouse. Fv/Fm decreased quadratically with increasing leaf temperature in a similar fashion in both tests, suggesting that the laboratory approach can adequately predict plant response to greenhouse heat stress.