Photosynthetic gas exchange and temperature-induced damage in seedlings of the tropical alpine species Argyroxiphium sandwicense

The capacity of Argyroxiphium sandwicense (silverword) seedlings to acclimate photosynthetic processes to different growing temperatures, as well as the tolerance of A. sandwicense to temperatures ranging from –15 to 60° C, were analyzed in a combination of field and laboratory studies. Altitudinal changes in temperature were also analyzed in order to explain the observed spatial distribution of A. sandwicense. A. sandwicense (Asteraceae) is a giant rosette plant that grows at high elevation on two Hawaiian volcanoes, where nocturnal subzero temperatures frequently occur. In addition, the soil temperatures at midday in the open alpine vegetation can exceed 60° C. In marked contrast to this large diurnal temperature variation, the seasonal variation in temperature is very small due to the tropical maritime location of the Hawaiian archipelago. Diurnal changes of soil and air temperature as well as photosynthetic photon flux density were measured on Haleakala volcano during four months. Seedlings were grown in the laboratory, from seeds collected in ten different A. sandwicense populations on Haleakala volcano, and maintained in growth chambers at 15/5, 25/15, and 30/25° C day/night temperatures. Irreversible tissue damage was determined by measuring electrolyte leakage of leaf samples. For seedlings maintained at each of the three different day/night temperatures, tissue damage occurred at –10° C due to freezing and at about 50° C due to high temperatures. Tissue damage occurred immediately after ice nucleation suggesting that A. sandwicense seedlings tend to avoid ice formation by permanent supercooling. Seedlings maintained at different day/night temperatures had similar maximum photosynthetic rates (5 mol m^–2 s^–1) and similar optimum temperatures for photosynthesis (about 16° C). Leaf dark respiration rates compared at identical temperatures, however, were substantially higher for seedlings maintained at low temperatures, but almost perfect homeostasis is observed when compared at their respective growing conditions. The lack of acclimation in terms of frost resistance and tolerance to high temperatures, as well as in terms of the optimum temperature for photosynthesis, may contribute to the restricted altitudinal range of A. sandwicense. The small seasonal temperature variations in the tropical environment where this species grows may have prevented the development of mechanisms for acclimation to longterm temperature changes.     

Molecular genetic variation following a population crash in the endangered Mauna Kea silversword, Argyroxiphium sandwicense ssp. sandwicense (Asteraceae)

The endangered Mauna Kea silversword, Argyroxiphium sandwicense ssp. sandwicense (Asteraceae), has experienced a severe decline in distribution and abundance because of predation by alien ungulates. The small remnant natural population on the Mauna Kea volcano contains only 46 individuals. By contrast, the Haleakala silversword, A. sandwicense ssp. macrocephalum, consists of a large, vigorous population exceeding 60 000 individuals. Molecular genetic variation in the two populations was assessed using random amplified polymorphic DNA (RAPD) loci. Despite its severe crash in size, the Mauna Kea population did not differ significantly from the Haleakala population in the number of detectably polymorphic loci or in heterozygosity. The lack of substantial reduction in genetic variation, at least as measured with RAPD loci, suggests that the Mauna Kea population may not yet have gone through multiple generations at very small size.     

Central alpha-MSH,energy balance,thermal balance,and antipyresis

Intracerebroventricular administration of alpha-MSH in young adult rats enhanced metabolic rate and caused a dose-dependent suppression of food intake, exhibiting a coordinated catabolic pattern. However, the thermoregulatory effects did not seem to be coordinated: the rising heat production was accompanied by a practically simultaneous tendency for rise in heat loss (skin vasodilatation), and the final core temperature either increased or decreased depending on which rise prevailed. The effect on heat loss possibly explains the antipyretic properties of the peptide.     

Responses to Mg/Ca balance in an Iranian serpentine endemic plant, Cleome heratensis (Capparaceae) and a related non-serpentine species, C. foliolosa

A soil Ca/Mg quotient greater than unity is generally considered necessary for normal plant growth but some serpentine plants are adapted to much lower Ca/Mg quotients, resulting from a major cation imbalance in their substrata. In order to investigate the growth and tolerance responses of serpentine and non-serpentine species to varied Ca/Mg quotients, controlled nutrient solution experiments were performed using an a newly reported Iranian endemic serpentine plant, Cleome heratensis Bunge et Bien. Ex Boiss. and a related non-serpentine species Cleome foliolosa DC. and a Eurasian Ni-hyperaccumulating species Alyssum murale Waldst. and Kit. Seedlings were grown in modified Hoagland’s solutions with varying Ca and Mg concentrations (0.2–2.5 and 0.5–10 mM, respectively) in a fully factorial randomised block design. The yields of the two serpentine plants increased significantly as Mg concentrations in the nutrient solution were increased from 0.5 to 4 mM but decreased in the 10 mM Mg treatment. For C. foliolosa yields decreased significantly from 0.5 to 10 mM Mg, indicating the sensitivity of this non-serpentine plant, and the relative tolerance of the serpentine plants to extremely high levels of Mg. Shoot and root Mg and Ca concentrations in C. heratensis and A. murale were higher than those in C. foliolosa in the low and moderate Mg treatments, supporting the view that many serpentine plants have a relatively high requirement for Mg. Maximum Mg concentrations were found in the roots of C. heratensis. Yields of C. heratensis and A. murale did not change significantly as Ca levels in nutrient solution increased from 0.2 to 2.5 mM Ca, However the yield of C. foliolosa increased significantly from 0.2 to 1.5 mM Ca, indicating sensitivity in this non-serpentine plant and tolerance of the two serpentine plants to low levels of Ca correlated with tissue Ca concentrations, probably because of a greater ability for Ca uptake at low-Ca availability. Calcium deficiency in the low-Ca treatments could be a reason for reduced yield in the non-serpentine plants.     

Influence of insulating dead leaves and low temperatures on water balance in an Andean giant rosette plant

Abstract. The influence of insulating dead leaves on water balance in Espeletia timotensis Cuatr., an Andean caulescent giant rosette plant, was studied under field and laboratory conditions. Removal of the dead leaf layer surrounding the stem changed the pattern of diurnal stem temperature variation and produced transient and permanent effects on water balance. The pattern of liquid water flow resistance increase at low stem temperatures suggested that much of the water flow in the stem was through living membranes, probably those of the pith cells. The pith was determined to be an important source of stored water for daily transpirational needs. The lethal effects of dead leaf removal were attributed to one or more of the following causes: (1) inhibition of pith recharge by subfreezing stem temperatures; (2) embolisms in stem xylem; (3) freezing injury to pith tissue. The results suggested that an insulating layer of marcescent leaves and the presence of an internal water reservoir closer to the rosette than the soil water are important adaptations for maintenance of a favourable water balance in tropical alpine habitats where freezing temperatures occur regularly but last only a few hours.     

Water use by the desert cucurbit Citrullus colocynthis (L.) Schrad.

Summary The rates of water use and leaf surface conductance of Citrullus colocynthis (Cucurbitacea) were evaluated from measurements of the surface temperature and microenvironment of leaves. At desert sites in Saudi Arabia the transpiration rates reached 0.13–0.17 g m^-2 s^-1 and the leaf temperatures were always close to air temperature. Leaf models (dry) placed in the canopy were considerably warmer than the air. To investigate responses over a wider range of conditions, plants were grown in a controlled environment room. It was found that when conditions were made hotter than those that occurred in the desert, the stomatal conductance increased greatly. Transpiration rate attained 0.6 g m^-2 s^-1 and the leaves were up to seven degrees cooler than the air. The results suggest a finely-tuned control mechanism working like a switch when the leaves experience extreme conditions, and enabling the plant to avoid lethal temperatures.     

The role of capacitance in the water balance of Andean giant rosette species

Abstract Pith water storage capacity and its role in plant-water relations were studied in seven giant rosette species of the genus Espeletia from the Venezuelan Andes. Readily available water from the pith was calculated to be capable of sustaining mean transpiration for up to 2.5 h. The relative importance of water stored in the pith, however, differed among species. The species that grow in the higher and colder environments tended to have a greater capacitance than the species that grow in the lower and less extreme environments. The pith volume per unit leaf area (PV/LA) was found to be a good indicator of the relative water storage capacity of the adult individuals of each species. Diurnal fluctuations in leaf water potential were not as pronounced in the species with higher PV/LA values. The species-specific PV/LA was highly correlated with the leaf turgor loss point and with the total resistance to water flow from soil to leaves. These results suggested that species-specific capacitance in the genus Espeletia is a response to temperature-limited soil water availability and that cold tropical environments with frequent subfreezing temperatures tend to select for high water storage capacity in giant rosette plants.     

Climate and the meristem temperatures of plant communities near the tree-line

Summary Temperatures of terminal meristems of forest, krummholz and dwarf shrub vegetation were measured at altitudes of 450, 600, 650 and 850 m in the Cairngorm Mountains of Scotland. Simultaneously, the air temperature above the vegetation was recorded, so that it was possible to calculate the difference between meristem and air temperature, sometimes called the excess temperature. This temperature increased linearly with the net radiation absorbed at each station, and the slope was dependent on wind speed and the height of the vegetation. In the extreme cases the slopes were practically zero for forest and 0.028° C W^-1 m² for dwarf shrubs. The latter implies a temperature excess of about 15° C in bright sunshine and low wind speeds. A model is developed to calculate the excess temperature from a knowledge of vegetational height and climatological variables.     

Heat-induced accumulation of chloroplast protein synthesis elongation factor, EF-Tu, in winter wheat

Chloroplast protein synthesis elongation factor, EF-Tu, has been implicated in heat tolerance in maize (Zea mays). Chloroplast EF-Tu is highly conserved, and it is possible that this protein may be of importance to heat tolerance in other species including wheat (Triticum aestivum). In this study, we assessed heat tolerance and determined the relative levels of EF-Tu in mature plants (at flowering stage) of 12 cultivars of winter wheat experiencing a 16-d-long heat treatment (36/30 degrees C, day/night temperature). In addition, we also investigated the expression of EF-Tu in young plants experiencing a short-term heat shock (4h at 43 degrees C). Heat tolerance was assessed by examining the stability of thylakoid membranes, measuring chlorophyll content, and assessing plant growth traits (shoot dry mass, plant height, tiller number, and ear number). In mature plants, relative levels of EF-Tu were determined after 7 d of heat stress. High temperature-induced accumulation of EF-Tu in mature plants of all cultivars, and a group of cultivars that showed greater accumulation of EF-Tu displayed better tolerance to heat stress. Young plants of all cultivars but one did not show significant increases in the relative levels of EF-Tu. The results of the study suggest that EF-Tu protein may play a role in heat tolerance in winter wheat.     

Invasive litter, not an invasive insectivore, determines invertebrate communities in Hawaiian forests

In Hawaii, invasive plants have the ability to alter litter-based food chains because they often have litter traits that differ from native species. Additionally, abundant invasive predators, especially those representing new trophic levels, can reduce prey. The relative importance of these two processes on the litter invertebrate community in Hawaii is important, because they could affect the large number of endemic and endangered invertebrates. We determined the relative importance of litter resources, represented by leaf litter of two trees, an invasive nitrogen-fixer, Falcataria moluccana, and a native tree, Metrosideros polymorpha, and predation of an invasive terrestrial frog, Eleutherodactylus coqui, on leaf litter invertebrate abundance and composition. Principle component analysis revealed that F. moluccana litter creates an invertebrate community that greatly differs from that found in M. polymorpha litter. We found that F. moluccana increased the abundance of non-native fragmenters (Amphipoda and Isopoda) by 400% and non-native predaceous ants (Hymenoptera: Formicidae) by 200%. E. coqui had less effect on the litter invertebrate community; it reduced microbivores by 40% in F. moluccana and non-native ants by 30% across litter types. E. coqui stomach contents were similar in abundance and composition in both litter treatments, despite dramatic differences in the invertebrate community. Additionally, our results suggest that invertebrate community differences between litter types did not cascade to influence E. coqui growth or survivorship. In conclusion, it appears that an invasive nitrogen-fixing tree species has a greater influence on litter invertebrate community abundance and composition than the invasive predator, E. coqui.     

Effects of manipulation of plant carbon nutrient balance on tall goldenrod resistance to a gallmaking herbivore

Summary Ramets from six Solidago altissima clones of known resistance to the stem gallmaker Eurosta solidaginis were grown with and without supplemental nutrients. In a greenhouse experiment, mated female Eurosta were allowed to oviposit in ramets that were subsequently grown through flowering and then harvested to determine biomass allocation. Supplemental nutrients increased plant biomass but did not affect resistance to this gallmaking herbivore. This result does not conform to the plant carbon/nutrient balance hypothesis which predicts that enhanced mineral nutrition will indirectly cause a reduction in carbon-based defensive chemistry. Our results indicate a strong genetic basis to ball gallmaker resistance since modification of host phenotype did not influence susceptibility. We suggest that evaluating the degree of genotypic or environmental control of plant resistance will be especially helpful in clarifying the patterns of defensive chemical responses.     

Salinity effect on plant growth and leaf demography of the mangrove, Avicennia germinans L.

We assessed the effect of salinity on plant growth and leaf expansion rates, as well as the leaf life span and the dynamics of leaf production and mortality in seedlings of Avicennia germinans L. grown at 0, 170, 430, 680, and 940 mol m⁻³ NaCl. The relative growth rates (RGR) after 27 weeks reached a maximum (10.4 mg g⁻¹ d⁻¹) in 170 mol m⁻³ NaCl and decreased by 47 and 44% in plants grown at 680 and 940 mol m⁻³ NaCl. The relative leaf expansion rate (RLER) was maximal at 170 mol m⁻³ NaCl (120 cm m⁻² d⁻¹) and decreased by 57 and 52% in plants grown at 680 and 940 mol m⁻³ NaCl, respectively. In the same manner as RGR and RLER, the leaf production (P) and leaf death (D) decreased in 81 and 67% when salinity increased from 170 to 940 mol m⁻³ NaCl, respectively. Since the decrease in P with salinity was more pronounced than the decrease in D, the net accumulation of leaves per plant decreased with salinity. Additionally, an evident increase in annual mortality rates (λ) and death probability was observed with salinity. Leaf half-life (t _0.5) was 425 days in plants grown at 0 mol m⁻³ NaCl, and decreased to 75 days at 940 mol m⁻³ NaCl. Thus, increasing salinity caused an increase in mortality rate whereas production of new leaves and leaf longevity decreased and, finally, the leaf area was reduced.     

A newScaevola (Goodeniaceae) from Kauai Hawaiian plant studies 92

Based on morphology, there is describedScaevola wahiawaensis, native to the mountains of Kauai Island, Hawaiian Islands. It has the blades tomentose below with erect, subequal hairs; corolla lateral lobes 12–14 mm long, 6–7 mm wide, and the thickened body 2.5–3 mm wide. Its closest relative,S. mollis, of Oahu, has the blades closely tomentose below and ascending pilose; corolla lateral lobes 7–11 mm long, 3–3.5 (−5) mm wide, and the thickened body 1.5 mm wide.     

Carbon and water balance in Polylepis sericea,a tropical treeline species

Polylepis sericea trees grow well above the continuous forest line in the Venezuelan Andes. In these environments, extreme daily temperature ranges can occur at any time of the year and trees experience a 4 month dry period. The purpose of this work was to study carbon and water relations of this species in the field during wet and dry seasons in order to understand this species' success at such high altitudes. Leaf gas exchange (portable system in open mode) and leaf water potential (pressure chamber) were measured at 1–2 h intervals during several daily courses at 4000 m elevation in the Páramo de Piedras Blancas. CO₂ assimilation versus leaf temperature curves were also obtained for this species in the laboratory. Clear differences in the measured parameters were observed between seasons. For a wet season day, maximum CO₂ assimilation rate was 7.4 mol m^-2 s^-1 and leaf conductance was relatively constant (approximately 100 mmol m^-2 s^-1)In the dry season day, maximum CO₂ assimilation rate was 5.8 molm^-2 s^-1 and leaf conductance was close to 60 mmolm^-2 s^-1. Minimum leaf water potentials measured were -1.3 MPa for the wet and -2.2 MPa for the dry season. The CO₂ assimilation-leaf temperature relationship showed a 13.4°C leaf temperature optimum for photosynthesis with maximum and minimum compensation points of 29.5 and -2.8°C, respectively. Maximum night-time respiration was relatively high (2.7 (imol) m^-2 s^-1)Our results show thatP. sericea maintains a highly positive carbon balance through all daily courses, even though there is a slight water stress effect during the dry season; this suggests that its carbon assimilation machinery is well adapted to the low temperatures and seasonal water stress found in the high tropical mountains.     

Adequacy of food rations in soldiers during exercise in hot, day-time conditions assessed by doubly labelled water and energy balance methods

The energy requirements of people doing physical work in hot climates are not clearly understood. In particular, we know little about the combined effects of heat stress and muscular work on energy requirements. During military exercises in the African bush soldiers are supplied with standard rations, the adequacy of which is unknown. We have now assessed the adequacy of these food and water rations in 12 male Zimbabwean soldiers during 12 days of strenuous, heat-stress exercise in the field. We used two methods to measure energy expenditure: the double-labelled water method (DLW) and the energy balance method (i.e. comparing dietary energy with changes, if any, in body energy stores). Two groups were studied: one group (eight subjects) carried out field exercises; the control group consisted of four soldiers doing normal work. Mean daily energy expenditure as assessed by the DLW method was [mean (SE)] 23 (1.5) MJ · day⁻¹ for the field group and 14 (0.5) MJ · day⁻¹ for the control group (P<0.001). By the energy balance method, daily energy expenditure was calculated to be 26 (0.7) MJ · day⁻¹ and 15.5 (0.4) MJ · day⁻¹ for the field group and control group, respectively. Body mass loss was 3 (0.1) kg [4.6 (0.3)% of body mass] for the field group, but the control group gained 1.1 (0.1) kg. Mean daily fluid intake was 11 (0.5) 1 · day⁻¹, suggesting that the standard ration supplied was inadequate. Body mass loss was caused by both the energy deficit and total body water loss. These results suggest strenuous work in hot, dry field conditions imposes extra energy requirements. Accepted: 21 January 1997     

Comparison of the physiology, morphology, and leaf demography of tropical saplings with different crown shapes

Branch architecture, leaf photosynthetic traits, and leaf demography were investigated in saplings of two woody species, Homolanthus caloneurus and Macaranga rostulata, co-occurring in the understory of a tropical mountain forest. M. rostulata saplings have cylindrical crowns, whereas H. caloneurus saplings have flat crowns. Saplings of the two species were found not to differ in area-based photosynthetic traits and in average light conditions in the understory of the studied site, but they do differ in internode length, leaf emergence rate, leaf lifespan, and total leaf area. Displayed leaf area of H. caloneurus saplings, which have the more rapid leaf emergence, was smaller than that of M. rostulata saplings, which have a longer leaf lifespan and larger total leaf area, although M. rostulata saplings showed a higher degree of leaf overlap. Short leaf lifespan and consequent small total leaf area would be linked to leaf overlap avoidance in the densely packed flat H. caloneurus crown. In contrast, M. rostulata saplings maintained a large total leaf area by producing leaves with a long leaf lifespan. In these understory saplings with a different crown architecture, we observed two contrasting adaptation strategies to shade which are achieved by adjusting a suite of morphological and leaf demographic characters. Each understory species has a suite of morphological traits and leaf demography specific to its architecture, thus attaining leaf overlap avoidance or large total leaf area.     

Geoecological alteration of surface soils by the Hawaiiansilversword (Argyroxiphium sandwicense DC.) inHaleakala's crater,Maui

The geoecological effects of organic matter addition to soil byHawaiian silverswords (Argyroxiphium sandwicense DC.,Asteraceae) in Haleakala's crater (Maui) are examined. Sets of 12 surfacesoil samples were gathered in three positions: bare soils; under the crowncanopy of adjacent live plants; and below dead, withered rosettes. Silverswordstanding litter and live foliage were also collected. Several physical andchemical properties were investigated. Bare soils were coarse and exceptionallylow in chemical nutrients and water-holding capacity. Litter addition belowsilverswords altered all properties except particle-size distribution; soilsunder dead rosettes became modified more strongly than those below live plants.While rosettes grow, standing litter is tightly preserved, thus only smallamounts of organic matter are added to soil. When plants die, a substantialinflux of litter is released in a short period (7 to 9 years average); thisraises the C and N content, C/N ratios, exchangeable nutrient levels (Ca, Mg,K, P) and cation exchange capacity of soil. Physical properties, such as color,bulk density and porosity are also greatly affected; as a result, soilwater-retention at field capacity nearly doubles under withering plants. Soiltemperatures (5 cm depth) at noon were 18.1°C lower beneath silversword canopies(18.5°C mean) than in contiguous bare soils(36.6°C). This cooling may result in lower evaporationand further conservation of soil moisture, thus increased seedgermination. Nutrient-retention strategies may have evolved in the semelparousArgyroxiphium to help it cope with the harsh,drought-prone soils and unpredictable climate of Haleakala. Silverswordflowering results in a tremendous seed output, but the plant needs to ensurethat some seeds – which have limited dispersal capability – willgerminate, and thus provides a nutrient-rich, water-retaining substrate moresuitable for seedling establishment than bare soils. Through itsself-regulating influence on underlying soils,Argyroxiphium affects both soil formation and its ownregeneration in Haleakala.     

Soil and plant water relations in a crested wheatgrass pasture: response to spring grazing by cattle

Summary Few field studies have attempted to relate effects of actual livestock grazing on soil and plant water status. The present study was initiated to determine the effects of periodic defoliations by cattle during spring on soil moisture and plant water status in a crested wheatgrass (Agropyron cristatum (L.) Gaertn. and A. desertorum (Fisch. ex Link) Schult.) pasture in central Utah. Soil moisture in the top 130 cm of the soil profile was depleted more rapidly in ungrazed plots than in grazed plots during spring and early summer. Soil moisture depletion was more rapid in grazed plots in one paddock after 1 July due to differential regrowth, but there was no difference in soil water depletion between plots in another paddock during the same period. This difference in soil water depletion between paddocks was related to a difference in date of grazing. Although more water had been extracted from the 60 cm to 130 cm depths in ungrazed plots by late September, cumulative soil moisture depletion over the entire 193 cm profile was similar in grazed and ungrazed plots. Prior to 1 July, grazing had no effect on predawn leaf water potentials as estimated by a pressure chamber technique; however, after 1 July, predawn leaf water potentials were lower for ungrazed plants. Midday leaf water potentials were lower for grazed plants before 1 July, but did not differ between grazed and ungrazed plants after 1 July. A 4- to 8-day difference in date of defoliation did not affect either predawn or midday leaf water potentials. The observed differences in water use patterns during spring and early-summer may be important in influencing growth and competitive interactions in crested wheatgrass communities that are subject to grazing by domestic livestock.     

Effects of temperature on photosynthesis of two morphologically contrasting plant species along an altitudinal gradient in the tropical high Andes

The effects of temperature on photosynthesis of a rosette plant growing at ground level, Acaena cylindrostachya R. et P., and an herb that grows 20–50 cm above ground level, Senecio formosus H.B.K., were studied along an altitudinal gradient in the Venezuelan Andes. These species were chosen in order to determine – in the field and in the laboratory – how differences in leaf temperature, determined by plant form and microenvironmental conditions, affect their photosynthetic capacity. CO₂ assimilation rates (A) for both species decreased with increasing altitude. For Acaena leaves at 2900 m, A reached maximum values above 9 μmol m⁻² s⁻¹, nearly twice as high as maximum A found at 3550 m (5.2) or at 4200 m (3.9). For Senecio leaves, maximum rates of CO₂ uptake were 7.5, 5.8 and 3.6 μmol m⁻² s⁻¹ for plants at 2900, 3550 and 4200 m, respectively. Net photosynthesis-leaf temperature relations showed differences in optimum temperature for photosynthesis (A _o.t.) for both species along the altitudinal gradient. Acaena showed similar A _o.t. for the two lower altitudes, with 19.1°C at 2900 m and 19.6°C at 3550 m, while it increased to 21.7°C at 4200 m. Maximum A for this species at each altitude was similar, between 5.5 and 6.0 μmol m⁻² s⁻¹. For the taller Senecio, A _o.t. was more closely related to air temperatures and decreased from 21.7°C at 2900 m, to 19.7°C at 3550 m and 15.5°C at 4200 m. In this species, maximum A was lower with increasing altitude (from 6.0 at 2900 m to 3.5 μmol m⁻² s⁻¹ at 4200 m). High temperature compensation points for Acaena were similar at the three altitudes, c. 35°C, but varied in Senecio from 37°C at 2900 m, to 39°C at 3550 m and 28°C at 4200 m. Our results show how photosynthetic characteristics change along the altitudinal gradient for two morphologically contrasting species influenced by soil or air temperatures. Received: 5 July 1997 / Accepted: 25 October 1997