Growth and leaf physiology of monkeyflowers with different altitude ranges

Every species is limited both geographically and ecologically to a subset of available habitats, yet for many species the causes of distribution limits are unknown. Temperature is thought to be one of the primary determinants of species distributions along latitudinal and altitudinal gradients. This study examined leaf physiology and plant performance under contrasting temperature regimes of sister species of monkeyflower, Mimulus cardinalis and Mimulus lewisii (Phrymaceae), that differ in altitude distribution to test the hypothesis that temperature is the primary determinant of differences in fitness versus altitude. Each species attained greatest aboveground biomass, net photosynthetic rate, and effective quantum yield of photosystem II when grown under temperatures characteristic of the altitudinal range center. Although both species exhibited greater stem length, stomatal conductance, and intercellular CO₂ concentration in hot than in cold temperatures, these traits showed much greater reductions under cold temperature for M. cardinalis than for M. lewisii. Survival of M. lewisii was also sensitive to temperature, showing a striking decrease in hot temperatures. Within each temperature regime, the species native to that temperature displayed greatest growth and leaf physiological capacity. Populations from the altitude range center and range margin of each species were used to examine population differentiation, but central and marginal populations did not differ in most growth or leaf physiological responses to temperature. This study provides evidence that M. cardinalis and M. lewisii differ in survival, growth, and leaf physiology under temperature regimes characterizing their contrasting low and high altitude range centers, and suggests that the species’ altitude range limits may arise, in part, due to metabolic limitations on growth that ultimately decrease survival and limit reproduction.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Temperature and photoperiodic control of developmental responses in climatic races of Mimulus

Two strongly differentiated climatic races of the Mimulus cardinalis-lewisiicomplex were grown at a variety of temperatures (3–27°C)and photoperiods (8 and 16 hr) under controlled environmentalconditions. M. cardinalis (the lowland race, 400 m) and M. lewisii(the sub-alpine race, 3200 m) were found to differ in theirphysiological responses to the varied environments in severalsignificant ways: 1) At 27°C (16 hrphotoperiod), M. lewisiisustained 100% mortality in contrast to the substantial growthand flowering of M. cardinalis under these conditions; 2) In8 hr photoperiods at all temperatures, there was little growthand no flowering in M. lewisii whereas there was considerablegrowth at all temperatures, and flowering at 23 and 27°Cin M. cardinalis; 3) At low temperatures (7–15°C),16 hr photoperiods, flowering occurred a week or two earlierin M. lewisii than in M. cardinalis. The lowland race has asignificantly wider temperature and photoperiodic tolerancethan has the sub-alpine race. Applications of gibberellic acidto rosette Mimulus plants under non-inductive conditions (15°C,8 hr photoperiod) promoted vigorous stem elongation withoutflowering. The application of steroids, other hormones and metaboliteshad no observable effects. ¹Present address: Faculty of Botany, Ohio State University,Columbus, Ohio, U.S.A. (Received March 16, 1970; )     

CHROMOSOMAL REARRANGEMENTS AND THE GENETICS OF REPRODUCTIVE BARRIERS IN MIMULUS (MONKEY FLOWERS)

Chromosomal rearrangements may directly cause hybrid sterility and can facilitate speciation by preserving local adaptation in the face of gene flow. We used comparative linkage mapping with shared gene‐based markers to identify potential chromosomal rearrangements between the sister monkeyflowers Mimulus lewisii and Mimulus cardinalis, which are textbook examples of ecological speciation. We then remapped quantitative trait loci (QTLs) for floral traits and flowering time (premating isolation) and hybrid sterility (postzygotic isolation). We identified three major regions of recombination suppression in the M. lewisii × M. cardinalis hybrid map compared to a relatively collinear Mimulus parishii × M. lewisii map, consistent with a reciprocal translocation and two inversions specific to M. cardinalis. These inferences were supported by targeted intraspecific mapping, which also implied a M. lewisii‐specific reciprocal translocation causing chromosomal pseudo‐linkage in both hybrid mapping populations. Floral QTLs mapped in this study, along with previously mapped adaptive QTLs, were clustered in putatively rearranged regions. All QTLs for male sterility, including two underdominant loci, mapped to regions of recombination suppression. We argue that chromosomal rearrangements may have played an important role in generating and consolidating barriers to gene flow as natural selection drove the dramatic ecological and morphological divergence of these species.     

Chromosomal rearrangements directly cause underdominant F1 pollen sterility in Mimulus lewisii–Mimulus cardinalis hybrids

Chromosomal rearrangements can contribute to the evolution of postzygotic reproductive isolation directly, by disrupting meiosis in F₁ hybrids, or indirectly, by suppressing recombination among genic incompatibilities. Because direct effects of rearrangements on fertility imply fitness costs during their spread, understanding the mechanism of F₁ hybrid sterility is integral to reconstructing the role(s) of rearrangements in speciation. In hybrids between monkeyflowers Mimulus cardinalis and Mimulus lewisii, rearrangements contain all quantitative trait loci (QTLs) for both premating barriers and pollen sterility, suggesting that they may have facilitated speciation in this model system. We used artificial chromosome doubling and comparative mapping to test whether heterozygous rearrangements directly cause underdominant male sterility in M. lewisii–M. cardinalis hybrids. Consistent with a direct chromosomal basis for hybrid sterility, synthetic tetraploid F₁s showed highly restored fertility (83.4% pollen fertility) relative to diploids F₁s (36.0%). Additional mapping with Mimulus parishii–M. cardinalis and M. parishii–M. lewisii hybrids demonstrated that underdominant male sterility is caused by one M. lewisii specific and one M. cardinalis specific reciprocal translocation, but that inversions had no direct effects on fertility. We discuss the importance of translocations as causes of reproductive isolation, and consider models for how underdominant rearrangements spread and fix despite intrinsic fitness costs.     

Floral volatile alleles can contribute to pollinator‐mediated reproductive isolation in monkeyflowers (Mimulus)

Pollinator‐mediated reproductive isolation is a major factor in driving the diversification of flowering plants. Studies of floral traits involved in reproductive isolation have focused nearly exclusively on visual signals, such as flower color. The role of less obvious signals, such as floral scent, has been studied only recently. In particular, the genetics of floral volatiles involved in mediating differential pollinator visitation remains unknown. The bumblebee‐pollinated Mimulus lewisii and hummingbird‐pollinated Mimulus cardinalis are a model system for studying reproductive isolation via pollinator preference. We have shown that these two species differ in three floral terpenoid volatiles – d ‐limonene, β‐myrcene, and E‐β‐ocimene – that are attractive to bumblebee pollinators. By genetic mapping and in vitro analysis of enzyme activity we demonstrate that these interspecific differences are consistent with allelic variation at two loci, LIMONENE‐MYRCENE SYNTHASE (LMS) and OCIMENE SYNTHASE (OS). Mimulus lewisii LMS (MlLMS) and OS (MlOS) are expressed most strongly in floral tissue in the last stages of floral development. Mimulus cardinalis LMS (McLMS) is weakly expressed and has a nonsense mutation in exon 3. Mimulus cardinalis OS (McOS) is expressed similarly to MlOS, but the encoded McOS enzyme produces no E‐β‐ocimene. Recapitulating the M. cardinalis phenotype by reducing the expression of MlLMS by RNA interference in transgenic M. lewisii produces no behavioral difference in pollinating bumblebees; however, reducing MlOS expression produces a 6% decrease in visitation. Allelic variation at the OCIMENE SYNTHASE locus is likely to contribute to differential pollinator visitation, and thus promote reproductive isolation between M. lewisii and M. cardinalis. OCIMENE SYNTHASE joins a growing list of ‘speciation genes’ (‘barrier genes’) in flowering plants.     

Selective oviposition by a leaf miner in response to temporal variation in abscission

Summary Responses to humidity of net photosynthesis and leaf conductance of single attached leaves were examined in populations of herbs from wet soil sites in Beltsville, Maryland and Davis, California, USA. Plants were grown in controlled environments under three conditions which differed in the magnitude of the day-night temperature difference and in daytime air saturation deficit. No population differences in response were found in Abutilon theophrasti. In Amaranthus hybridus stomatal conductance and net photosynthesis were more reduced by increasing leaf to air water vapor pressure difference (VPD) in the population from Beltsville, but only for the growth condition with a constant 25°C temperature. In Chenopodium album, stomatal conductance was more sensitive to VPD in the population from Davis, but only for the growth condition with 28/22°C day/night temperatures. Population differences in the sensitivity to VPD of leaf conductance were associated with differences in leaf area to root weight ratio. The relative reduction of net photosynthesis as VPD increased was greater than, equal to, or less than the relative decrease in substomatal carbon dioxide partial pressure. The pattern depended on species, and on growth condition. From these results one can not conclude that environmental humidity has been a strong selective force in determining sensitivity to humidity of stomatal conductance.     

Leaf hairs: Effects on physiological activity and adaptive value to a desert shrub

Summary The effects of leaf hairs on photosynthesis, transpiration, and leaf energy balance were measured on the desert shrub Encelia farinosa in order to determine the adaptive significance of the hairs. The pubescence reduces leaf absorptance resulting in a reduced heat load, and as a consequence lower leaf temperatures and lower transpiration rates. In its native habitat where air temperatures often exceed 40° C, the optimum temperature for photosynthesis in E. farinosa occurs at 25° C, and at leaf temperatures above 35° C net photosynthesis declines precipitously. An advantage of leaf pubescence is that it allows a leaf temperature much lower than air temperature. As a result, leaf temperatures are near the temperature optimum for photosynthesis and high, potentially lethal leaf temperatures are avoided. However, there is a disadvantage associated with leaf pubescence. By reflecting quanta that might otherwise be used in photosynthesis, the presence of leaf hairs reduces the rate of photosynthesis. A tradeoff model was used to assess the overall advantage of possessing leaf hairs. In terms of the carbon gaining capacity of the leaf, the model predicted that for different environmental conditions different levels of leaf pubescence were optimal. In other words, under aird conditions and/or high air temperatures, leaves of E. farinosa would have a higher rate of photosynthesis by being pubescent than by not being pubescent. The predictions from this model agreed closely with observed patterns of leaf pubescence in the field.C.I.W.-D.P.B. Publication No. 613     

Effects of day-to-day changes in root temperature on leaf conductance to water vapour and CO2 assimilation rates of Vigna unguiculata L. Walp.

Summary Leaf gas exchange of Vigna unguiculata was influenced by short-term (day-to-day) changes in soil temperature and the response depended upon the aerial environment. When aerial conditions were constant at 30° C leaf temperature, high air humidity and moderate quantum flux, CO₂ assimilation rate and leaf conductance increased with increases in soil temperature from 20 to 35° C, and this response was reversible. Decreases in CO₂ assimilation rate and leaf conductance were observed at root temperatures above 30° C when root temperatures were increased from 20° C to 40° C and when air humidity was decreased in steps during the day. In contrast, varying soil temperatures between 20 to 35° C had no influence on gas exchange when shoots were subjected to a wide range of temperatures during each day.The gain ratio A/E remained constant at different air humidities when root temperature was less than or equal to 30° C indicating optimal gas exchange regulation, but changed with humidity at higher root temperatures. Leaf conductance responded independently from leaf water potential which remained relatively constant during individual experiments.The results indicate that plant responses to high root temperatures may have relevance to plant performance in semi-arid environments. They also illustrate the importance of controlling soil temperatures when studying the responses of potted plants in controlled aerial environments.Dedicated to K.F. Springer     

Influence of root temperature on growth of Pinus sylvestris,Fagus sylvatica,Tilia cordata and Quercus robur

One-year-old tree seedlings were incubated in a greenhouse from April to July, under natural daylight conditions, with their root systems at constant temperatures of 5, 10, 15, 20, 25, 30 and 35 °C and with the above ground parts kept at a constant air temperature of 18–20 °C. The course of height growth, total mass increment, root, shoot and leaf weight as well as leaf areas were measured. The results indicate that clear differences exist in the optimal root zone temperatures for various growth parameters in different tree species. Pinus sylvestris had a maximal height increment at about 5–10 °C and maximal total mass increment at 15 °C root temperature. In contrast, the optimum for Quercus robur was at 25 °C. Tilia cordata and Fagus sylvatica had their optima for most growth parameters at 20 °C. The root temperature apparently indirectly influenced photosynthesis (dry weight accumulation) and respiration loss. From the observed symptoms and indications in the literature it seems probable that a change in hormone levels is involved as the main factor in the described effects. Variation of root temperature had only an insignificant effect on bud burst and the time at which the shoots sprouted. Apparently species of northern origin seem to have lower root temperature optima than those of more southern origin. This is to be verified by investigation of other tree species.     

Influence of different soil temperatures on water use,growth and internal water status of soybean

Studies on the effect of soil temperature on internal water relations of well watered soybean (Glycine max L.) at constant air temperature under controlled conditions were carried out. A specially designed thermogradient tank was used for obtaining a range of soil temperatures. Data on shoot height, shoot weight, root length, root weight, leaf area and leaf water potential were obtained at 41 days after sowing and the highest values of these parameters were recorded at 28.6°C. The air temperature during the course of these investigations was 25±1°C and it may be concluded that slightly warmer soil temperatures than air temperatures were optimal for soybean with regard to the above measured parameters.     

Influence of cold soil and snowcover on photosynthesis and leaf conductance in two Rocky Mountain conifers

Summary The influence of cold soil and snowcover on photosynthesis and conductance of Picea engelmannii and Pinus contorta was investigated early in the growing season in the Medicine Bow Mountains, Wyoming, USA. Trees of both species growing in cold soil (<1°C) associated with snowpack had 25–40% lower leaf photosynthesis than trees in warm soils (>10°C). In cold soils leaf conductance of both species was lower, but more so in Pinus, leading to lower intercellular CO₂ concentrations and greater stomatal limitation of photosynthesis. Soil temperature had no effect on predawn and midday shoot water potentials of Pinus and Picea and lower photosynthesis and conductance did not appear to be a result of lower bulk shoot water potential. Predawn, as well as midday, water potentials of Pinus were consistently higher than Picea suggesting that Pinus may have deeper roots, although trenching experiments indicated young Picea trees have more extensive lateral root systems than similar sized Pinus trees. Young Picea trees (<2 m in height) in snowbanks were capable of utilizing warmer soil 4 m from their base. Under similar conditions Pinus in snowbanks had lower photosynthesis and conductance than controls and Pinus did not appear capable of utilizing warmer soils nearby. Under full sunlight, PPFD reflected from the snow surface was 400–1400 mol m^-2 s^-1 higher than from snow-free surfaces. This reflected light resulted in a 10%–20% increase in photosynthesis of Picea. The beneficial effect of reflected light was apparent whether or not photosynthesis was reduced by low soil temperatures.     

The influence of water stress on the photosynthetic performance and stomatal behaviour of tree seedlings subjected to variation in temperature and irradiance

Summary Seedlings of Betula pendula Roth. and Gmelina arborea L. were subjected to variation in temperature and irradiance. The influence of a mild water-stressing treatment on the photosynthetic performance and stomatal behaviour of these plants was assessed. For both species, the shape of the relationships between irradiance and photosynthesis and temperature and photosynthesis resembled those reported for other species. The effect of water stress was to reduce the rate of photosynthesis, particularly at high temperatures. This was largely a function of a reduction in mesophyll conductance under these conditions. The optimum temperature for stomatal opening was significantly lower than the optimum temperature for photosynthesis, which was in turn lowered by the water stress treatment. The stomata of birch seedlings showed maximum opening at an intermediate temperature while the stomata of Gmelina generally exhibited a closing movement when leaf temperatures increased from 15° C. Mesophyll conductances of both species increased with increasing temperature.The physiological basis for the variation in photosynthetic performance and stomatal behaviour and the ecological significance of this variation are discussed.     

Plant transpiration at high elevations: Theory,field measurements,and comparisons with desert plants

Summary The influence of elevational changes on plant transpiration was evaluated using leaf energy balance equations and well-known elevational changes in the physical parameters that influence water vapor diffusion. Simulated transpirational fluxes for large leaves with low and high stomatal resistances to water vapor diffusion were compared to small leaves with identical stomatal resistances at elevations ranging from sea level to 4 km. The specific influence of various air temperature lapse rates was also tested. Validation of the simulated results was accomplished by comparing actual field measurements taken at a low elevation (300 m) desert site with similar measurements for a high elevation (2,560 m) mountain research site. Close agreement was observed between predicted and measured values of transpiration for the environmental and leaf parameters tested.Substantial increases in solar irradiation and the diffusion coefficient for water vapor in air (D _wv) occurred with increasing elevation, while air and leaf temperatures, the water vapor concentration difference between the leaf and air, longwave irradiation, and the thermal conductivity coefficient for heat in air decreased with increasing elevation. These changes resulted in temperatures for sunlit leaves that were further above air temperature at higher elevations, especially for large leaves. For large leaves with low stomatal resistances, transpirational fluxes for low-elevation desert plants were close to those predicted for high-elevation plants even though the sunlit leaf temperatures of these mountain plants were over 10°C cooler. Simulating conditions with a low air temperature lapse rate (0.003° C m^-1 and 0.004° C m^-1) resulted in predicted transpirational fluxes that were greater than those calculated for the desert site. Transpiration for smaller leaves decreased with elevation for all lapse rates tested (0.003° C m^-1 to 0.010° C m^-1). However, transpirational fluxes at higher elevations were considerably greater than expected for all leaves, especially larger leaves, due to the strong influence of increased solar heating and a greater D _wv. These results are discussed in terms of similarities in leaf structure and plant habit observed among low-elevation desert plants and high-elevation alpine and subalpine plants.     

The temperature optima for tree seedling photosynthesis and growth depend on water inputs

Understanding how tree growth is affected by rising temperature is a key to predicting the fate of forests in future warmer climates. Increasing temperature has direct effects on plant physiology, but there are also indirect effects of increased water limitation because evaporative demand increases with temperature in many systems. In this study, we experimentally resolved the direct and indirect effects of temperature on the response of growth and photosynthesis of the widely distributed species Eucalyptus tereticornis. We grew E. tereticornis in an array of six growth temperatures from 18 to 35.5°C, spanning the climatic distribution of the species, with two watering treatments: (a) water inputs increasing with temperature to match plant demand at all temperatures (W_incr), isolating the direct effect of temperature; and (b) water inputs constant for all temperatures, matching demand for coolest grown plants (W_const), such that water limitation increased with growth temperature. We found that constant water inputs resulted in a reduction of temperature optima for both photosynthesis and growth by ~3°C compared to increasing water inputs. Water limitation particularly reduced the total amount of leaf area displayed at T_opt and intermediate growth temperatures. The reduction in photosynthesis could be attributed to lower leaf water potential and consequent stomatal closure. The reduction in growth was a result of decreased photosynthesis, reduced total leaf area display and a reduction in specific leaf area. Water availability had no effect on the response of stem and root respiration to warming, but we observed lower leaf respiration rates under constant water inputs compared to increasing water inputs at higher growth temperatures. Overall, this study demonstrates that the indirect effect of increasing water limitation strongly modifies the potential response of tree growth to rising global temperatures.     

Microhabitat segregation and physiological differences in co-occurring tiger beetle species,Cicindela oregona and Cicindela tranquebarica

Summary The daily movements of two co-occurring tiger beetle species were monitored in conjunction with changes in microclimate along streams in Northeast Arizona. Cicindela oregona and C. tranquebarica temporarily segregated across areas of beach exhibiting different microclimates. C. oregona progressively moved from the dry upper beach to the wet stream edge as beach temperatures increased and humidity decreased. The actively foraged throughout the day in this moist habitat at air temperatures between 25 and 38°C. C. tranquebarica remained on the dry, upper portions of the beach and shuttled between sun and shade at air temperatures above 35°C. Only when stream edge temperatures exceeded 30°C was tranquebarica found in this subhabitat. Both species exhibited physiological tolerances in the laboratory that were consistent with their microhabitat preferences in the field. Although both species had similar high lethal temperatures (47–48°C) in saturated air, oregona died at lower temperatures (39–43°C) than tranquebarica (46–47°C) under dry (0% RH) conditions. C. oregona was considerably more active than tranquebarica at body temperatures below 30°C and exhibited higher levels of active metabolism between 25 and 40°C. In addition, C. tranquebarica exhibited significantly lower water loss rates than oregona at 30, 35 and 40°C.     

Oxygen inhibition of photosynthesis

The effect of leaf temperature, O₂ and calculated O₂/CO₂ solubility ratio in the leaf on the quantum yield of photosynthesis was studied for the C₄ species, Zea mays L., and the C₃ species, Triticum aestivum L. Over a range of leaf temperatures of 16 to 35° C, the quantum yield of Z. mays was relatively constant and was similar under 1.5 and 21% O₂, being ca. 0.059 mol CO₂ mol^-1 quanta absorbed. Under 1.5% O₂ and atmospheric levels of CO₂, the quantum yield of T. aestivum was relatively constant (0.083 mol CO₂ mol^-1 quanta absorbed) at leaf temperatures from 15 to 35° C. Atmospheric levels of O₂ (21%) reduced the quantum yield of photosynthesis in T. aestivum and as leaf temperature increased, the quantum yield decreased from 0.062 at 15°C to 0.046 mol CO₂ mol^-1 quanta absorbed at 35°C. Increasing temperature decreases the solubility of CO₂ relatively more than the solubility of O₂, resulting in an increased solubility ratio of O₂/CO₂. Experimentally manipulating the atmospheric levels of O₂ or CO₂ to maintain a near-constant solubility ratio of O₂/CO₂ at varying leaf temperatures largely prevented the temperature-dependent decrease in quantum yield in t. aestivum. Thus, the decrease in quantum yield with increasing leaf temperature in C₃ species may be largely caused by a temperaturedependent change in the solubility ratio of O₂/CO₂.J and II=Ku and Edwards, 1977a, b     

CO2 and water vapour exchange in four alpine herbs at two altitudes and under varying light and temperature conditions

CO₂ and water vapour exchange rates of four alpine herbs namely: Rheum emodi, R. moorcroftianum, Megacarpaea polyandra and Rumex nepalensis were studied under field conditions at 3600 m (natural habitat) and 550 m altitudes. The effect of light and temperature on CO₂ and water vapour exchange was studied in the plants grown at lower altitude. In R. moorcroftianum and R. nepalensis, the average photosynthesis rates were found to be about three times higher at 550 m as compared to that under their natural habitat. However, in M. polyandra, the CO₂ exchange rates were two times higher at 3600 m than at 550 m but in R. emodi, there were virtually no differences at the two altitudes. These results indicate the variations in the CO₂ exchange rates are species specific. The change in growth altitude does not affect this process uniformly.The transpiration rates in R. emodi and M. polyandra were found to be very high at 3600 m compared to 550 m and are attributed to overall higher stomatal conductance in plants of these species, grown at higher altitude. The mid-day closure of stomata and therefore, restriction of transpirational losses of water were observed in all the species at 550 m altitude. In addition to the effect of temperature and relative humidity, the data also indicate some endogenous rhythmic control of stomatal conductance.The temperature optima for photosynthesis was close to 30°C in M. polyandra and around 20°C in the rest of the three species. High temperature and high light intensity, as well as low temperature and high light intensity, adversely affect the net rate of photosynthesis in these species.Both light compensation point and dark respiration rate increased with increasing temperature.The effect of light was more prominent on photosynthesis than the effect of temperature, however, on transpiration the effect of temperature was more prominent than the effect of light intensity.No definite trends were found in stomatal conductance with respect to light and temperature. Generally, the stomatal conductance was highest at 20°C.The study reveals that all these species can easily be cultivated at relatively lower altitudes. However, proper agronomical methodology will need to be developed for better yields.     

Influence of krummholz mat microclimate on needle physiology and survival

Summary Microclimate and photosynthesis of krummholz mat growth forms of Picea engelmanii (Parry) and Abies lasiocarpa [Hook.] Nutt. were investigated to determine structural features which may aid survival in alpine environments. The structure of krummholz mats was described in terms of the vertical distribution of leaf area index and leaf area density, which exceeded 50 m^-1 (based on total leaf surface area) near the canopy surface and approached zero below 30 cm from the surface in both species. Photosynthetic photon flux density (PPFD, 0.4–0.7 m wavelengths) and wind decreased by an average of 6 and 50-fold, respectively, between 1 m above and 10 cm below mat surfaces in both species. Needle temperatures on a P. engelmannii krummholz mat during July averaged about 2°C above air temperature during the day, with a maximum overtemperature of greater than 20°C above T _air during one sunlit period. At night, needle temperatures averaged 3–4°C below T _air.Net photosynthesis in year-old P. engelmannii shoots reached a maximum at 15–20°C during July and August. Surface shoots were light saturated at near 1200 moles m^-2s^-1 PPFD, and had higher photosynthetic rates than subsurface, predominantly shaded shoots above 800 moles m^-2s^-1. Shade shoots had higher photosynthetic rates when PPFD was below 600 moles m^-2s^-1, and at 250 moles m^-2s^-1 shade shoots maintained about 50% of the net photosynthetic rate of sun shoots at light saturation. Shade shoots appeared capable of benefitting photosynthetically from elevated temperatures within krummholz mats despite relatively low light levels. Especially rapid photosynthesis may occur when canopy needles are illuminated by sunflecks and needle temperatures rise by 10° C or more.Snow cover appears crucial for the survival of needles during winter. Snow accumulated within krummholz needle canopies before the sub-canopy zone of unfoliated branches became filled. The concentrated needle growth in the krummholz canopy captured snow in early autumn without support from ground-level snowpack. Early snow cover in both species prevented cuticle abrasion and resulted in high winter needle water contents and viabilities for subsurface compared to surface needles which became abraded, severely dehydrated, and had high mortality between December and February, especially on windward sides of shoots.Extremely high concentrations of needles within krummholz mat canopies created an aerodynamic structure which elevated needle temperatures to more optimal photosynthetic levels in summer and resulted in more efficient snow accumulation in winter. These factors appear crucial for winter needle survival. Thus, krummholz mats appear to be an important adaptation in growth form which provides survival benefits in both summer and winter.     

Leaf temperatures and energy balance ofWelwitschia mirabilis in its natural habitat

Summary Welwitschia mirabilis is a perennial desert plant with extremely large leaves (0.5–1.0 m broad, 1–2 m long). Leaf temperatures were measured in the field and the energy budget was calculated. The portions of the leaf which were kept above the ground had leaf temperatures which were only 4–6°C above air temperature. In the leaf portions which were in contact with the ground leaf temperatures were 6–12°C above air temperature (absolute maximum 51°C). The important feature in the energy budget ofWelwitschia mirabilis is its high reflectivity (38% of the global radiation). Only about 56% of the global radiation is absorbed by the thick leathery leaves. The energy loss due to convection is of the same order of magnitude as the reflection and it is abouy the same in the portions of leaf on and above the ground. The difference in leaf temperatures found in these portions is due to the loss of thermal radiation from the section of leaf above the ground to the cooler ground which is shaded by the leaf. The provision of a heat sink due to the large area of shade cast by these large leaves is of significance to the existence ofWelwitschia mirabilis in its arid habitats.     

FLAVONOID PIGMENTS IN MIMULUS CARDINALIS AND ITS RELATED SPECIES. I. ANTHOCYANINS

Chromatographic and spectrophotometric techniques were used to identify the anthocyanin pigments present in Mimulus cardinalis and its related species of section Erythranthe of the genus Mimulus (Scrophulariaceae). On the basis of rigorous tests, the flowers of M. cardinalis were found to contain pelargonidin-3-glucoside, pelargonidin-3-rhamnoglucoside, the caffeoyl ester of pelargonidin-3-glucoside, cyanidin-3-glucoside, cyanidin-3-rhamnoglucoside, and the caffeoyl ester of cyanidin-3-glucoside. Qualitatively all members of the group contain these six anthocyanins except M. lewisii. All of its populations lack the pelargonidin glycosides and some lack, in addition, the cyanidin-3-rhamnoglucoside. The striking differences in flower color and intensity appear to be due to quantitative differences not here analyzed.