Temperature and high salinity effects in germinating dimorphic seeds of Atriplex rosea

Atriplex rosea L. (Chenopodiaceae; tumbling orach), an annual herb, is a widely established weedy species of disturbed sites in all counties of Utah. Seeds of Atriplex rosea were collected from a salt marsh in Faust, Utah, and are dimorphic, light brown, and 2-2.5 mm wide, or black and 1-2 mm wide. Seed germination responses of the black and brown seeds were studied over a range of salinity and temperature. Both brown and black seeds germinated at 1000 mM NaCl, and the optimal temperature for germination of both types was 20°-30°C. Variation in temperature, however, affected germination of black seeds more than brown seeds. At lower thermoperiod only 40%-50% black seeds germinated in nonsaline control, and germination was almost completely inhibited with the inclusion of salinity. However, all brown seeds germinated in control at temperatures above 5°-15°C, and inhibition caused by salinity was comparatively lower. Brown seeds had a higher germination rate than black seeds at all temperature and salinity treatments. The highest rate of germination of both seeds occurred at the temperature regime of 5°-15°C. Recovery of germination for black seeds when transferred to distilled water after being in various salinity treatments for 20 days was quite variable. Recovery decreased with increase in salinity at lower temperature regimes, increased with salinity at optimal thermoperiod, and had no effect at 20°-30°C. Brown seeds recovered poorly from salinity at all thermoperiods except 5°-15°C, where recovery decreased with an increase in salinity. Brown seeds are adapted to germination in the early part of the growing season, whereas black seeds are capable of surviving harsher conditions and can germinate in later time periods. Characteristics of the dimorphic seeds increase chances for survival in the harsh saline desert environment.     

Effects of osmotic potential, potassium chloride, and sodium chloride on germination of greasewood ( Sarcobatus vermiculatus )

Greasewood ( Sarcobatus vermiculatus [Hook.] Torr.) (Chenopodiaceae) typically grows on salt-affected soils where its germination requirements may reflect characteristics necessary for establishment in saline environments. The objective of this study was to determine the effect of osmotic potential and specific ions on the germination of seeds from three populations of greasewood. Seeds were germinated at 20 C in solutions of polyethylene glycol with water potentials ranging from –0.3 to –2.2 MPa that contained 0 to 68480 µmol·L –1 sodium chloride (NaCl) or 0 to 53640 µmol·L –1 potassium chloride (KCl). Germination of two populations was reduced by increasing salt concentration and decreasing osmotic potential; germination of one population was reduced by declining osmotic potential. No seeds germinated at an osmotic potential lower than –1.6 MPa. For all populations, days to 50% of final germination increased and abnormal germination decreased as osmotic potential declined. Comparison of our results with those from other studies suggests geographic ecotypic development in response to osmotic potential and NaCl and KCl concentrations during germination.     

High rates of photosynthesis in the desert shrub Chrysothamnus nauseosus ssp. albicaulis

  Basic aspects of photosynthesis were investigated in white rubber rabbitbrush ( Chrysothamnus nauseosus (Pallas) Britt. ssp. albicaulis ), a common C 3 deciduous shrub native to arid regions of the western U.S. Under favorable field conditions, net photosynthesis (P n ) ranged from 36 to 73 mgCO 2 · dm – 2 · hr – 1 , which is relatively high for a woody species. The leaves from the actively growing flowering shoots exhibited higher P n than those on the vegetative shoots. P n also varied according to the age of the leaves and the location of the plants. P n did not light saturate even at quantum flux densities (QFD) equivalent to full sunlight. The light compensation point was relatively high (ca 100 μ mol · m – 2 · S – 1 ), perhaps due to the presence of a tomentose vestiture on the leaf surface. At high QFD's, the stomatal conductance was high (ca 520 mmol · m – 2 · s – 1 ) for a woody species. RUBP - carboxylase content of the leaves ranged from 20 to 22 mg per gram F.W., which is similar to that found in most C 3 crop species. These results suggest that rabbitbrush is able to maintain high rates of P n , at least under nonstressed conditions.           

Chloroplast ultrastructure in the desert shrub Chrysothamnus nauseosus ssp. albicaulis

Ultrastructure of the chloroplasts of white rubber rabbitbrush ( Chrysothamnus nauseosus (Pallas) Britt. ssp. albicaulis ) was observed with electron microscopy. In addition, leaf anatomy was observed with light microscopy. Previously, it had been reported that the leaves of this desert shrub exhibited a relatively high rate of photosynthesis when compared to other C 3 plants. Comparisons with chloroplasts of other C 3 and C 4 plants demonstrated a reduced amount of granal stacking in the rabbitbrush. However, the classification of rabbitbrush as a C 3 plant is confirmed. RUBP-carboxylase concentration is reported at about 450 mg · ml -1 stromal space based on the estimation of 1 mg of chlorophyll per 25 ul of stromal space in a normal C 3 chloroplast and data from an assay to determine the ratio of RUBP-carboxylase to chlorophyll.     

Effect of salinity on seed germination of Triglochin maritima under various temperature regimes

Triglochin maritima L. (arrow grass), an herbaceous perennial in the family Juncaginacea, is widely distributed in inland and coastal salt marshes of North America. Triglochin maritima seeds from a population growing in a salt marsh at Faust, Utah, were germinated at 4 temperature regimes (12-h night/12-h day, 5-15° C, 10-20° C, and 15-25° C) and 5 salinities (0, 100, 200, 300, 400, and 500 mol m -3 NaCl) to determine optimal conditions for germination and level of salt tolerance. Ungerminated seeds were returned to distilled water after 20 d to determine whether seeds could recover from salinity treatments. Maximum germination occurred in distilled water, and increases in NaCl concentration progressively inhibited seed germination. No seeds germinated at concentrations higher than 400 mol m -3 NaCl. A temperature regime of low night (5° C) and high day (25° C) temperature yielded maximum germination; all other temperature regimes significantly inhibited seed germination relative to this optimum. Recovery of germination was highest at 5-25° C and lowest at 5-15° C. Recovery of seed germination when seeds were transferred to distilled water from salt solutions was highest at 5-25° C (72%) for seeds exposed to the 500 mol m -3 NaCl pretreatment and significantly reduced at other temperature regimes. The recovery germination response indicates a synergistic inhibitory interaction effect on germination when seeds were exposed to high salinities at suboptimal thermoperiods.     

Chrysothamnus nauseosus ssp. uintahensis: a stabilized hybrid

Chrysothamnus nauseosus ssp. uintahensis (Asteraceae) is formally described and illustrated. The new subspecies represents stabilized hybrids that most probably have C. nauseosus ssp. hololeucus x C. parryi ssp. attenuatus ancestry. Morphological and anatomical comparisons are made among the putatively related taxa.        

Improving seed germination of Salicornia rubra (Chenopodiaceae) under saline conditions using germination-regulating chemicals

Salicornia rubra Nels. (Chenopodiaceae) is a salt-tolerant annual species occurring in salt playas of the Great Basin desert in western United States. It forms pioneer communities on the most saline location of an inland salt playa at Goshen, Utah. Seeds of Salicornia rubra are capable of germinating at 1000 mM NaCl at 25-35° C alternating temperature regime. Dormancy-regulating chemicals were evaluated for their ability to alleviate the innate and salinityenforced dormancy in seeds of Salicornia rubra . Betaine, GA3, kinetin, fusicoccin, ethephon, thiourea, proline, and nitrate had no effect in alleviating primary dormancy. Increases in NaCl concentration progressively inhibited germination of Salicornia rubra seeds. Ethephon, fusicoccin, GA3, kinetin, thiourea, and nitrate promoted germination under low saline conditions. At high salinity fusicoccin had no effect, whereas GA3, kinetin, and ethephon substantially alleviated salinity effects. Application of these dormancy-regulating compounds could be of practical value in seeding a saline area for restoration purposes, particularly under high saline conditions.     

Variation in germination response to temperature and water availability in blackbrush ( Coleogyne ramosissima ) and its ecological significance

Blackbursh ( Coleogyne ramosissima Torr.) is a dominant desert shrub in a distinct mid-elevations vegetation belt between creosote bush-bursage ( Larrea tridentata-Ambrosia dumosa ) shrubland below and big sagebrush-pinyon pine- Utah juniper ( Artemisia tridentata-Pinus monophylla-Juniperus osteosperma ) woodland above in the Mojave Desert. Seed germination patterns of blackbrush seeds collected from 2 elevations (1200 and 1550 m) in 5 mountain ranges within the blackbrush shrublands were investigated. Morphological features of blackbrush seeds, including weight, length, and width, were not significantly different ( P > 0.05) among elevations and mountain ranges in the Mojave Desert. Germination of blackbrush seeds was optimal when preceded by a prechill period of 4-6 wk. Seeds incubated at room temperature germinated poorly. Seeds collected at warm, low-elevation sites appeared to be less dormant (required less prechill time), germinated faster, and showed a higher overall germination response; watering at 2-wk intervals revealed the greatest germination. Some ecotypic variation among populations establishing at different elevations was evident with regard to dormancy duration and germination response at certain constant temperatures.     

Chrysothamnus nauseosus ssp. iridis (Asteraceae): a new endemic from Utah

Chrysothamnus nauseosus ssp. uintahensis (Asteraceae) is formally described and illustrated. The new subspecies represents stabilized hybrids that most probably have C. nauseosus ssp. hololeucus x C. parryi ssp. attenuatus ancestry. Morphological and anatomical comparisons are made among the putatively related taxa.        

Subspecies specificity of gall forms on Chrysothamnus nauseosus

Galls induced by tephritid flies (Diptera: Tephritidae) on Chrysothamnus nauseosus ssp. albicaulis, consimilis, graveolens, and salicifolius are useful in taxonomic identification of these subspecies in Utah. In this study, “callus” galls were found almost exclusively on ssp. albicaulis. “Cotton” galls were more frequent on ssp. consimilis and graveolens. “Mace” galls occurred on all four subspecies but were the only gall found on naturally occurring populations of ssp. salicifolius. Gall form specificity is not as well correlated with C. nauseosus subspecies in peripheral areas of the species range as in Utah. This decrease in specificity may be due to decreased competition for the host plant in outlying areas.     

Desiccation sensitivity and heat tolerance of Prunus ilicifolia seeds dispersed by American black bears ( Ursus americanus )

Carnivore consumption of fruit is a principle means by which many fleshy-fruited plant species achieve long-distance seed dispersal. We examined carnivore dispersal of hollyleaf cherry ( Prunus ilicifolia ) seeds, specifically assessing the survival, desiccation sensitivity, and germination of seeds found in bear scats. Studies were conducted both in the laboratory and in 2 burn areas in Los Padres National Forest, California. Bear scats containing P. ilicifolia seeds were collected in burned and unburned chaparral. We counted seeds in each scat and noted whether endocarps had tooth punctures or rattled audibly when shaken. For comparative germination trials, we also collected fruits and seeds directly from mature shrubs. In the laboratory, following a cold-moist stratification period, seeds were assessed for germinability. In the field, we compared desiccation rates and germinability of seeds from bear scats and freshly collected seeds. We compared rates of moisture loss and germination for seeds subjected to several different conditions, including 25 °C (room temperature), 30 °C and 65 °C (in the lab), and placement on exposed soil in a burn area (in the field), where midday temperatures were approximately 45 °C but likely much higher by late afternoon. Prunus ilicifolia seeds collected from bear scats were largely undamaged; the vast majority of these seeds germinated. In some cases, germination rate was higher for seeds from scats than for seeds from intact fruits. Several results indicate that desiccation is an important cause of reduced germinability. First, seeds that rattled audibly germinated poorly; and the louder the rattle, the lower the germination percentage. Second, seeds (both fresh and those from bear scats) placed in the field under protective screens had greatly reduced levels of germination (an 84% decline) after only 7 days. Third, seeds dried in the lab, even at relatively moderate temperatures, showed a decline in germination with seed moisture loss. The addition of high temperatures accelerated this decline in germination. We discuss the relevance of heat and desiccation sensitivity of seeds dispersed by bears to successful seed germination in burned and late-seral mesic and xeric chaparral. El consumo de fruta por carnívoros es uno de los principales medios por los que muchas especies de plantas de frutos carnosos logran dispersar sus semillas a través largas distancias. Examinamos la dispersión por carnívoros de las semillas del islay ( Prunus ilicifolia ), un arbusto del chaparral, evaluando concretamente la supervivencia, la sensibilidad a la desecación y la germinación de semillas encontradas en las heces de oso. Estos estudios se llevaron a cabo tanto en el laboratorio como en dos áreas quemadas en Los Padres National Forest, California. Se colectaron heces de oso que contenían semillas de P. ilicifolia en el chaparral quemado y en el no quemado. Contamos las semillas en cada bola fecal y notamos si los endocarpios tenían marcas de dientes y si vibraban audiblemente al agitarse. Para llevar a cabo pruebas comparativas de germinación, también colectamos frutos y semillas directamente de los arbustos maduros. En el laboratorio, después de un período de estratificación fría-húmeda, evaluamos la capacidad de germinación de las semillas. En el campo, comparamos las tasas de desecación y de germinación de semillas de las heces de oso con aquellas de semillas colectadas directamente de los arbustos. Se compararon las tasas de pérdida de humedad y germinación de las semillas sujetas a diversas condiciones, entre ellas 25 °C (temperatura ambiente), 30 °C y 65 °C en el laboratorio, y en el campo sobre suelo expuesto en un área quemada, donde la temperatura al mediodía era ~45 °C pero probablemente mucho más alta por la tarde. En general, las semillas de P. ilicifolia colectadas en las heces de oso no estaban dañadas; la gran mayoría de estas semillas germinaron; en algunos casos la tasa de germinación fue más alta que la de frutos intactos. Varios resultados indicaron que la desecación es una causa importante de la reducción en la capacidad de germinación. Primero, las semillas que vibraban audiblemente germinaron poco, y cuanto más fuerte el sonido, menor la germinación. Segundo, las semillas (tanto las frescas como las colectadas de heces de oso) colocadas en el campo bajo una malla protectora exhibieron niveles de germinación muy reducidos (una disminución del 84%) después de sólo siete días. Tercero, la germinación de las semillas desecadas en el laboratorio, incluso a temperaturas relativamente moderadas, disminuyó con la pérdida de humedad de las semillas. Temperaturas más altas aceleraron esta disminución en la germinación. Discutimos la importancia de la sensibilidad al calor y a la desecación en las semillas dispersadas por los osos para la germinación exitosa en el chaparral mésico y árido, tanto en el chaparral recién quemado como en el que está en las etapas finales de sucesión.     

Effects of forest fuel smoke on dwarf mistletoe seed germination

Seeds of three species of dwarf mistletoe, Arceuthobium americanum Nutt. Ex Engelm., A. cyanocarpum Coulter & Nelson, and A. vaginatum subsp. cryptopodum (Engelm.) Hawksw. & Wiens, were exposed to smoke from burning forest fuels. Premeasured amounts of coniferous needles and branch wood were burned in a small incinerator with smoke passing through a closed chamber containing the seeds. Following three different smoke treatments and one high-temperature treatment, tests were conducted to evaluate the effects of these treatments on seed germination. Germination was inhibited for all species when the seeds were exposed to smoke for 60 minutes or longer. Seeds of A. americanum were unaffected by exposures to smoke from drier fuels. The percentage of germinating seeds of A. cyanocarpum and A. vaginatum showed little effect from exposures of up to 30 minutes.     

Recovery and germinability of native seed fed to cattle

Using livestock as seed dispersal agents may be an effective method for increasing species diversity on degraded and previously seeded rangelands. We quantified seed passage and recovery rates, and post-passage germinability of Wyoming big sagebrush ( Artemisia tridentata Nutt. ssp. wyomingensis Beetle & Young), bottlebrush squirreltail ( Elymus elymoides [Raf.] Swezey), and gooseberry globemallow ( Sphaeralcea grossulariaefolia [H. & A.] Rydb.) by feeding Holstein heifers seeds of each species at 3 levels (15,000; 30,000; and 60,000 seeds) over a period of 3 weeks. One-kg fecal samples were collected 1, 2, 3, and 4 days after seed ingestion. Undamaged seeds were extracted from the samples and tested for germinability. Globemallow had the highest percentage of recovered, undamaged seed, followed by squirreltail and sagebrush. Globemallow and sagebrush seed passage was highest on Day 1, after which seed numbers dropped sharply. Squirreltail passage and recovery were more consistent through time, with higher seed recovery at lower seed feeding levels. Post-passage germinability was highest for squirreltail and globemallow on Day 1. Sagebrush germination was negligible. Differences in physical seed properties (size, shape, and seed coat) likely influenced interspecies variation in passage, recovery, and germinability. Globemallow and squirreltail seeds may be suited for livestock dispersal, but sagebrush seeds are not.     

Interspecific differences in grass seed imbibition

Seeds from 12 grass species were studied relative to mode of wetting and time of exposure to water to document interspecific differences in imbibition characteristics. Imbibition causes seeds to become wet, and wet seeds are more detectable to consumers than dry seeds. Thus, germination potential and ability to remain undetected by consumers may represent an important trade-off. Seeds wetted for 0-192 h in vials imbibed water at rates equivalent to seeds wetted by contact with wet paper towels pressing against their seed coat, except for seeds of Avena sativa , which weighed more after wetting in 2-mL vials with free water (0.471 g vs. 0.432 g). Seeds from different species imbibed water at different rates. These data show that interspecific variation in imbibition for seeds is high and support an expectation that imbibition potential can interact with detectability to consumers in an evolutionary trade-off.     

A sagebrush wilt disease of unknown cause

In a rare 1969 epidemic, spruce cone rust caused by Chrysomyxa pirolata infected 40 – 100 percent of trees and 20 – 67 percent of cones on riparian Colorado blue spruce on plots located in a 2200 – 2400 m elevational zone in Huntington Canyon of central Utah. Uredinial and telial sporulation on Pyrola spp. began in mid - June, a time closely correlated with opening of pistillate spruce cones. Come phenology and host habitat, as influenced by elevation, are apparently important factors in the restricted niche of the cone rust fungus in Utah. Several preceding consecutive years with extended periods of spring and fall moisture were associated with occurrence of the epidemic, although no cause - and - effect relationship was established. Weather records indicate that these events are infrequent in this climatic zone, and there was no detectable recurrence of cone rust for at least 9 years following 1969. Outwardly normal seeds developed in diseased cones, but seed germinability was reduced by 25 percent. Aecial spore masses between cone scales, cone resinosis, and distortion of cone scales prevented seed dispersal to the extent that the seed crop was effectually destroyed.     

Occurence and effect of Chrysomyxa pirolata cone rust on Picea pungens in Utah

In a rare 1969 epidemic, spruce cone rust caused by Chrysomyxa pirolata infected 40 – 100 percent of trees and 20 – 67 percent of cones on riparian Colorado blue spruce on plots located in a 2200 – 2400 m elevational zone in Huntington Canyon of central Utah. Uredinial and telial sporulation on Pyrola spp. began in mid - June, a time closely correlated with opening of pistillate spruce cones. Come phenology and host habitat, as influenced by elevation, are apparently important factors in the restricted niche of the cone rust fungus in Utah. Several preceding consecutive years with extended periods of spring and fall moisture were associated with occurrence of the epidemic, although no cause - and - effect relationship was established. Weather records indicate that these events are infrequent in this climatic zone, and there was no detectable recurrence of cone rust for at least 9 years following 1969. Outwardly normal seeds developed in diseased cones, but seed germinability was reduced by 25 percent. Aecial spore masses between cone scales, cone resinosis, and distortion of cone scales prevented seed dispersal to the extent that the seed crop was effectually destroyed.     

Respiration characteristics differ among cheatgrass ( Bromus tectorum L.) populations

Cheatgrass ( Bromus tectorum L.) is a dominant weed that has increased the frequency of wildfire in the Great Basin since its introduction about 100 yr ago. This study examines characteristics of respiratory metabolism in several different populations. Seeds from 6 populations were germinated and metabolic heat rates (q) and dark respiration rates (R co2 ) of all seedlings were measured calorimetrically at 15° and 25° C or (for 3 populations) at 5° intervals from 5° to 35° C. Growth rates, substrate carbon conversion efficiencies, and Arrhenius temperature coefficients were calculated from the data. Results show that cheatgrass metabolism is most efficient at temperatures near 0° C; at temperatures above 20-25° C, efficiency goes to zero. Cheatgrass populations differ in their temperature dependencies of substrate carbon conversion efficiency and predicted growth rate. Measurements of respiratory heat and CO 2 rates as functions of temperature can be made relatively quickly and used to aid understanding of metabolic adaptation by invasive and native species to the environment.     

Temperature-mediated changes in seed dormancy and light requirement for Penstemon palmeri (Scrophulariaceae)

Penstemon palmeri is a short-lived perennial herb colonizing disturbed sites in semiarid habitats in the western USA. In this study seed was harvested from six native and four seeded populations during two consecutive years. In laboratory germination trials at constant 15 C, considerable between-lot variation in primary dormancy and light requirement was observed. Four weeks of moist chilling (1 C) induced secondary dormancy at 15 C. Cold-induced secondary dormancy was reversed by one week of dark incubation at 30 C. This warm incubation treatment also reduced the light requirement of unchilled, after-ripened seed. Fluctuations in dormancy and light requirement of buried seeds have been linked to seasonal changes in soil temperature. Penstemon palmeri germination responses to temperature appear to be similar to those of facultative winter annuals.     

Feeding behavior and performance of a rabbitbrush leaf-beetle ( Trirhabda lewisii ) feeding on Chrysothamnus nauseosus regrwoth after fire

Fire often positively affects the growth and nutrient content of plants regrowing after a burn. These changes have been associated with preferential feeding by herbivores in burned areas. In this study in southeastern Wyoming, Chrysothamnus nauseosus Pursh (rubber rabbitbrush) regrowing after a fire produced new shoots with a distinct growth form. Shoots were longer than those on unburned control sites and had longer leaves with longer internodes between leaves. We conducted feeding trials to detect whether C. nauseosus shoots regrowing after fire were nutritionally superior to shoots from unburned plants for the specialist leaf beetle, Trirhabda lewisii Crotch (Coleoptera: Chrysomelidae). We also measured C:N ratios and nitrogen and water contents of leaves from burned and unburned plants. Trirhabda lewisii adults preferred shoots from burned plants when given a choice. The beetles ate similar amounts of burned and unburned plants when fed only a single type. Females that were fed either burned or unburned plants did not differ in number of eggs laid. Chemical analyses revealed no significant differences in nitrogen or water content of leaves from burned versus unburned plants. Carbon-to-nitrogen ratio of burned plants was marginally lower compared with unburned plants. In contrast to previous studies, which suggest that herbivore attraction to burned areas leads to enhanced performance, our study shows that performance is not necessarily enhanced after fire.     

Tamarix aphylla : a newly invasive tree in southern Nevada

In the southwestern United States, the nonnative athel pine ( Tamarix aphylla ) was presumed to be sterile and therefore not as likely to spread as its widely distributed, nonnative congener, T. ramosissima . However, at Lake Mead National Recreation Area (LMNRA) in southern Nevada, populations of T. aphylla have recently spread beyond their limited pre-1990 distribution and now form extensive monospecific stands. Over a 3-year period, we quantified seed production and germination from 60 T. aphylla trees at LMNRA. The annual mean seed production period was 50.6 days, and the mean potential germination (under laboratory conditions) was 22%, indicating that T. aphylla trees at LMNRA are capable of sexual reproduction in southern Nevada. No seeds germinated in field experiments, apparently because of high soil salt levels. However, seedling regeneration is becoming increasingly common at LMNRA. Tamarix aphylla trees occupied a distinct zone along the shoreline of Lake Mead, above T. ramosissima in elevation and below native Larrea tridentata communities, suggesting either competitive exclusion or differential resource utilization. The T. aphylla zone tended to have the highest mean values for total vegetation cover, leaf litter depth, soil salinity, soil moisture, pH, total Kjeldahl nitrogen, and soil organic matter. The capacity for sexual reproduction of this alien plant, combined with a suite of characteristics shared with the invasive T. ramosissima (e.g., drought tolerance and copious saline leaf litter), makes T. aphylla a potentially invasive species along the shores of LMNRA and other mesic areas in the desert Southwest.