Lichens are one of the common dominant biota in biological soil crusts (biocrusts), a community that is one of the largest in extent in the world. Here we present a summary of the main features of the lifestyle of soil crust lichens, emphasizing their habitat, ecophysiology and versatility. The soil crust is exposed to full light, often to high temperatures and has an additional water source, the soil beneath the lichens. However, despite the open nature of the habitat the lichens are active under shady and cooler conditions and avoid climate extremes of high temperature and light. In temperate and alpine habitats they can also be active for long periods, several months in some cases. They show a mixture of physiological constancy (e.g. similar activity periods and net photosynthetic rates) but also adaptations to the habitat (e.g. the response of net photosynthesis to thallus water content can differ for the same lichen species in Europe and the USA and some species show extensive rhizomorph development). Despite recent increased research, aspects of soil crust ecology, for example under snow, remain little understood.

Two field experiments were conducted to examine the competitive effect of common cocklebur and sicklepod in soybean. Factors examined included sap flow of both the weed and the crop, as well as the traditional parameters of weed and crop growth and crop yield. The intent was to determine to what extent competitive effects could be attributed to differential water utilization. In the first study, soybean planted at 20 plants m−1 row was infested with either sicklepod at 10 plants m−1 row or common cocklebur at two plants m−1 row. Sicklepod reduced soybean sap flow and yield 53 and 49%, respectively. Common cocklebur reduced soybean sap flow and yield 31 and 38%, respectively. Within a weed species, weed-induced yield and sap flow reductions were comparable, indicating that water deprivation was the primary detriment that these weeds inflicted on soybean. Individual common cocklebur plants were 3.9 times more competitive than sicklepod with respect to reducing soybean yield. However, sap flow of individual common cocklebur plants was only 1.4 times more than that of sicklepod, indicating that water deprivation was not the only causal factor in weed-crop competition. In the second experiment, sicklepod was planted in soybean (fixed density) and clipped to either half the height, same height as soybean, or left unclipped, in an attempt to simulate herbicide-induced stunting. Clipping sicklepod had a positive effect on soybean sap flow and yield. Thus, the benefits of reduced weed competition were achieved without killing sicklepod.

The influence of water stress on johnsongrass [Sorghum halepense (L.) Pers. ♯ SORHA] physiology was evaluated in a semiarid environment. Stomatal conductance of johnsongrass responded to more negative leaf water potential and increasing leaf temperature. The sensitivity of the leaf temperature effect was dependent on the soil water content. At low soil water content, conductance was limited by low water potential, and increasing leaf temperature had little effect. Conductance of CO2 was related to net photosynthesis in a curvilinear manner, with conductance levels greater than 0.3 mol·m-2· s-1 being in excess of that necessary for maximum photosynthesis. At both high conductance levels and low levels associated with increased water stress, intercellular CO2 concentration increased, indicating nonstomatal limitations to photosynthesis. Decreased osmotic potential provided the highest correlation with the linear decline of photosynthetic rate as stress intensified. The expression of osmotic adjustment in johnsongrass is reported during grain filling. Plants in the milkdough stage of grain filling had approximately 0.3 MPa lower osmotic potential at any relative water content than those at anthesis.

The effects of caffeic acid, chlorogenic acid, t-cinnamic acid, p-coumaric acid, ferulic acid, gallic acid, p-hydroxybenzaldehyde, 5-sulfosalicylic acid, vanillic acid, and vanillin on growth, photosynthesis, water relations, and chlorophyll content of 3-week-old soybeans [Glycine max (L.) Merr. ‘Tracy’] grown in aerated nutrient solution were determined. At concentrations of 10−3 M, caffeic, t-cinnamic, p-coumaric, ferulic, gallic, and vanillic acids significantly reduced dry matter production, leaf expansion, height, leaf production, net assimilation rate (rate of dry matter production per unit leaf area), and leaf area duration (total leaf area present during treatment interval). Chlorogenic acid, p-hydroxybenzaldehyde, 5-sulfosalicylic acid, and vanillin at 10−3 M did not inhibit growth. None of the 10 compounds at 10−4 M inhibited growth. At concentrations of 10−3 M, caffeic, t-cinnamic, p-coumaric, ferulic, gallic, and vanillic acids severely reduced net photosynthetic rate and stomatal conductance of single, fully expanded leaves. These same compounds also caused marked reductions in leaf chlorophyll content, with net losses of chlorophyll occurring over an 86-h period after treatment.

Sweet corn (Zea mays L. rugosa 'Seneca Scout’) and soybeans (Glycine max L. Merr. ‘Kanrich’) were grown in nutrient solutions containing alachlor [2-chloro-2′,6-diethyl-N-(methoxymethyl)acetanilide] at 2.36 × 10-5 M, polyethylene glycol MW 6000 (PEG6000) at −2.5 bars osmotic potential, alachlor at 2.36 × 10-5 M plus PEG6000 at −2.5 bars osmotic potential or with no additions. Sweet corn and soybean roots exposed to alachlor plus PEG6000 accumulated significantly more PEG6000 than roots exposed to PEG6000 alone; increased shoot uptake occurred only in soybean. Alachlor was also shown to significantly increase root osmotic potential and decrease root electrolyte leakage.

Increased osmotic potential from 0 to −14 bars decreased the moisture uptake and germination of mechanically scarified hemp sesbania [Sesbania exaltata (Raf.) Cory] seed. Germination percentage for osmotic potential of 0, −2, −4, −6, and −8 bars was 86%, 86%, 70%, 19%, and 1%, respectively. This response to moisture stress was modified by imbibition, or hydration-dehydration, of the seed prior to the moisture stress. Imbibing the seed for as little as 2 h significantly increased its subsequent germination against moisture stress. One hydration-dehydration cycle had little or no effect on germination, but as the number of cycles increased, the germination percentage decreased. Prolonged hydration of 3 and 7 days followed by dehydration greatly reduced subsequent seed germination.

Greenhouse and growth chamber experiments with potted plants were conducted to determine the effects of interspecific root and canopy interference, light intensity, and soil moisture on water uptake and biomass of soybean, common cocklebur, and sicklepod. Canopy interference and canopy plus root interference of soybean with common cocklebur increased soybean water uptake per plant and per unit leaf area. Root interference with soybean decreased common cocklebur water uptake per plant. Canopy interference of soybean with sicklepod increased soybean water uptake per unit leaf area, while root interference decreased uptake per plant. Combined root and canopy interference with soybean decreased water uptake per plant for sicklepod. Soybean leaf area and shoot weight were reduced by root interference with both weeds. Common cocklebur and sicklepod leaf area and shoot weight were reduced by root and canopy interference with soybeans. Only common cocklebur root weight decreased when canopies interfered and roots did not. The relationship between light intensity and water uptake per unit leaf area was linear in both years with water uptake proportional to light intensity. In 1991 water uptake response to tight was greater for common cocklebur than for sicklepod. The relationship between soil moisture level and water uptake was logarithmic. Common cocklebur water uptake was two times that of soybean or sicklepod at −2 kPa of pressure potential. In 1991 common cocklebur water uptake decreased at a greater rate than soybean or sicklepod in response to pressure potential changes from −2 to −100 kPa.

In a 6-yr field study, johnsongrass growing in ‘Forrest’ soybeans for the entire growing season reduced yields 45% in the year with most rainfall and 69% in the year with least rainfall. Yields of soybeans grown without johnsongrass competition were 56% higher in the year with the most rainfall than in the year with least rainfall. Yields of soybeans with full-season competition were reduced 75% from those in the year with most rainfall as compared with those in the year with least rainfall. The number of weeks of johnsongrass competition required to significantly reduce soybean yields varied from 3 wk, in the year with least rainfall, to 8 wk, in the year with most rainfall. Nine applications of sethoxydim at 0.84 kg ai ha-1 or haloxyfop at 0.55 kg ai ha-1 each year did not reduce soybean growth or yields.

The objective of this study was to fit a hydrothermal germination model to germination data for a seedlot of radiata pine (Pinus radiata D. Don). Seeds were incubated for 50 d at constant temperatures and water potentials (T = 12.5–32.5°C, Ψ = 0 to − 1.2 MPa). Most seeds completed germination within 50 d, but for low Ψ and/or non-optimal temperatures (T < 17.5°C, T>25°C) many seeds did not complete germination. In general, germination data conformed to the hydrothermal model. Departures from the model were encountered for slow-germinating seeds at suboptimal temperatures (T ≤ 20°C). To account for these departures, two alternative hydrothermal models were fitted with an additional term for an upwards shift in seed base water potential with increasing time to germination. The alternative models more correctly predicted germination time than the original model. Similarly, reduced percentage germination at supra-optimal temperatures (T>20°C) was explained by including a term in the hydrothermal model which shifted the base water potential of seeds upwards towards zero, which in turn reduced the predicted rate that hydrothermal time would be accumulated by seeds. The rate of this upwards shift in base water potential was dependent on time to complete germination and ambient water potential as well as supra-optimal temperature.

Patterns of edge-related environmental changes and plant water relations were investigated in the isolated forest reserves of the INPA-WWF Minimum Critical Size of Eco-systems project near Manaus, Brazil early in the wet season.

Air temperature was elevated and humidity reduced in the understorey within 40 m of the reserve margins, and air temperature and vapour pressure deficit (VPD) were higher in the interiors of 1 ha reserves than in 100 ha reserves. There was increased photosynthetically active radiation penetration to understorey level up to 40 m into a 100 ha reserve.

Soil moisture was depleted in the outer 20 m of both small and large reserves, and surface soil water potentials fell below – 1.5 MPa at the margin of a 1 ha reserve.

Studies of leaf relative water contents (RWCs) in understorey shrubs revealed no appreciable saturation deficits, though RWCs were sometimes lower at the reserve margins. Studies of leaf conductances revealed no evidence of restriction of water loss in these plants, and conductances of plants near the edges were significantly higher and higher for longer.

The implications of these results for reserve design and for local and regional water budgets, as well as the possible role of water stress in increased tree mortality in isolated reserves are discussed.

Padrōes de mudanças ambientais e hídricos de plantas relacionados a bordas de matas isoladas foram investigadas no início da estação seca nas reservas de floresta do Projeto Dinâmica Biológica de Fragmentos Florestais do INPA-WWF, proximo de Manaus, Brasil.

No sub-bosque, até 40 m da borda da mata, a temperatura ambiental estava elevada e a umidade reduzida, e a temperatura do ar e o DPV estavam maiores nas reservas de 1 ha do que nas reservas de 100 ha. A penetração de radiaçao fotosintéticamente ativa no sub-bosque estava aumentada até 40 m da borda em reservas de 100 ha.

A umidade do solo baixou consideravelmente nos 20 m periféricos de reservas grandes a pequenas, e o potencial hídrico na superficie do solo baixou para menos de −1.5 MPa na margem de uma reserva de 1 ha.

Estudos do teor relativo de água (TRA) em folhas de arbustos de sub-bosque nӑo revelaram deficits de saturação apreciaveis apesar de que TRAs foram menores nas bordas das reservas. Estudos da conductancia foliar não revelaram evidência de restrições de perdas hídricas nessas plantas, e as conductancias de plantas próximas a bordas foram significativamente maiores durante um período mais longo.

As impliçõtes desses resultados para o planejamento de reservas e para o balanço hídrico local e regional, assim como as possiveis funçoes de ‘stress’ hídrico na alta mortalidade de árvores em reservas isoladas são discutidas.

One component of the high organismal diversity of tropical lowland rain forests is the existence of series of closely related, sympatric species. For example, the six distinct tree species of the monophyletic Polyalthia hypoleuca complex regularly grow sympatrically in various combinations throughout the rain forests of Malesia. Theoretical and empirical evidence has been presented indicating that the members of such series either (1) are well differentiated with respect to at least one major niche component; or (2) may lack differentiated niches. In this study, certain sympatric members of the complex are shown to have clearly diverged in autecological characteristics that affect their distribution including: (1) P. glauca, P. hypoleuca, and P. sumatrana differ in growth characteristics, height at maturity, seedling germination requirements, and are found on soils with differing degrees of hydration; (2) P. discolor grows to a different height and on different substrate types than does P. multinervis; and (3) P. glauca and P. hypoleuca seedlings differ in transpiration characteristics and response to flooding and drought.

Lettuce (Lactuca sativa L.) seeds are inhibited from germinating above an upper temperature limit that is dependent upon cultivar, growing conditions and seed treatments. Thermoinhibition is accompanied by an increasing sensitivity of germination to reduced water potentials (ψ). We have employed a water relations analysis (hydrotime model) of seed germination rates to investigate the basis of thermoinhibition. Germination rates can be characterized by the distribution of base water potentials among seeds in the population (ψb(g)) and a hydrotime constant indicating the total accumulated hydrotime (MPa · h) above the base required for radicle emergence. The hydrotime model adequately described germination time courses across a range of ψ at both high and low temperatures. Increasing temperature caused the ψb(g) distributions to become more positive, accounting for the greater sensitivity to ψ and inhibition of germination. Increases in embryo osmotic potential and in the turgor yield thresholds of both the radicle and the endosperm/pericarp envelope contributed to this change. Seed priming (prehydration and drying) treatments speeded germination by reducing the hydrotime requirement. Priming also resulted in smaller increases in ψb(g) at high temperature, alleviating thermoinhibition by lowering the embryo yield threshold sufficiently to compensate for the increased endosperm resistance. The beneficial effects of priming in lettuce appear to occur primarily in the embryo, rather than in the surrounding envelope tissues.

A water relations (hydrotime) model was employed to characterize the responses of lettuce (Lactuca sativa L.) seed germination to temperature, water potential (ψ), endosperm disruption, and the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC). The base water potentials (ψb), standard deviations of base water potential and hydrotime constants (θH) estimated by the model were used to predict the germination rates at reduced ψ and supraoptimal temperatures in the presence or absence of ACC. The distribution of ψb values among seeds in the population (ψb(g)) increased (became more positive) as the imbibition temperature increased, indicating a greater sensitivity of seeds to reduced ψ at thermoinhibitory temperatures. Slitting the seeds at the cotyledonary end to disrupt the integrity of the endosperm extended the high temperature range for germination and maintained lower ψb(g) distributions compared with intact seeds as temperature increased. ACC (10 mm) in the imbibition solution raised the upper temperature limit for germination by about 2°C, and also lowered ψb(g) values by 0.1 to 0.2 MPa. The effect of ACC on ψb(g) was more pronounced at higher imbibition temperatures and in slit seeds. ACC (via conversion to ethylene) extends the high temperature limit for lettuce seed germination by acting in the embryo to maintain a lower ψ threshold for the initiation of growth as temperature increases. Evidence from other species as well suggests that this may be a general mechanism for the action of ethylene in promoting seed germination.

Embryogenesis was induced on cotyledons of immature zygotic embryos of soybean (Glycine max (L.) Merrill) placed on solid medium containing 62.5 mm glutamine, soybean seed growth medium salts and vitamins, and 40 mg I−1 2,4-dichlorophenoxyacetic acid (2,4-D) plus 175 mm maltose, or 8 mg I−1 α-naphthaleneacetic acid (NAA) plus 88 mM sucrose. Somatic embryo development was continued in liquid medium containing 0.16 mg I−1 indole-3-butyric acid and 2.64 mg I−1 abscisic acid, glutamine and salts as above, and 88–438 mM sucrose in progressively increasing steps. Germination was on solid half-strength Murashige-Skoog medium. During maturation, somatic embryos mimicked zygotic embryos in colour, protein concentration, water and solute potentials, and respiration. Protein and lipid accumulated to 329 and 86 g kg−1 dry weight in somatic embryos. Fatty acid composition was similar to that of axes of mature seeds. Before desiccation, the water and solute potentials of maturing somatic embryos declined to −1.13 and −1.99 MPa while turgor increased to 0.86 MPa. Concomitantly, a 60% reduction in activity of the cytochrome oxidase pathway of respiration occurred with somatic embryo maturation at 600 g water kg−1 fresh weight. Although small (about 8 mg per embryo), 60% of the somatic embryos formed roots and shoots during germination after maturation without drying and 30% germinated after drying to 60 g water kg−1 fresh weight. In the greenhouse, somatic plantlets grew to mature plants with seeds.

Quercus robur L. fruits are desiccation sensitive and shed from the plant at high moisture content (c. 48%). Water relations measurements were taken at intervals during the germination of prematurely-harvested immature fruits and naturally-shed mature fruits. In fruits from both harvests the pericarp and seed coat delayed germination by providing physical barriers to emergence of the radicle and by restricting the rate of imbibition. Although immature fruits could germinate in the absence of water they germinated more rapidly in the presence of an external water supply. The fully-mature fruit required a supply of external water, but germination was more rapid than in immature fruits. Substantial physiological changes, resulting in the accumulation of solutes, and increased axis size and extensibility occurred before splitting of the pericarp and subsequent germination of immature fruits. Such changes did not occur in fully-mature fruits, but fruits, from both harvests imbibed water and splitting of the pericarp appeared to result from increased embryo size and tissue pressure. Splitting was minimal prior to germination in immature fruits without an external water supply. It is suggested that the resulting reduction in Ψp when the pericarp splits was greater in the axis than in the cotyledons because of its greater capacity for expansion. This would create a Ψw gradient driving water flow to the axis from the cotyledons allowing continued radicle growth for germination.

Rainfall and soil moisture availability vary greatly both spatially and temporally. They are prime factors influencing species distribution patterns, diversity and habitat associations in a range of biomes, and limit primary productivity in many natural ecosystems, as well as in forestry and agricultural systems (Hawkins et al. 2003, Kozlowski & Pallardy 1997, Lieth 1975). Projections of drying trends, and increased frequency and intensity of drought events with climate and land-use changes (Hulme & Viner 1998, IPCC 2001) have fuelled an increased interest in the role of drought in determining the structure and function of natural and managed forest systems (Allen & Breshears 1998, Miles et al. 2004). Such projections accentuate the need to assess, understand and predict plant reactions to drought, as well as soil moisture variation at different scales.

Drought strongly influences plant phenology, growth and mortality in tropical forests, thereby shaping plant performance, population dynamics and community structure (Bunker & Carson 2005, Condit et al. 1995). Microbial symbionts of plants profoundly influence host water relations (Lösch & Gansert 2002), but are rarely considered in studies of tropical plant physiology. In particular, plant–fungus associations, which are ubiquitous in plant communities and especially common in tropical forests, play important and varied roles in plant water status. Fungal pathogens associated with roots, vascular tissue and foliage may interfere with water uptake and transport, increase rates of foliar transpiration, and induce xylem embolism and tissue death (Agrios 1997). In contrast, rhizosphere mutualists such as ecto- and arbuscular mycorrhizal fungi may benefit hosts by increasing surface area for water uptake, enhancing stomatal regulation of water loss, and increasing root hydraulic conductivity (Auge 2001, Lösch & Gansert 2002).

Dew deposition was calculated for leaves of the epiphytic bromeliads Tillandsia brachycaulos and T. elongata in a tropical dry deciduous forest in Yucatán, Mexico, over one year. Plant characteristics and water relations were also measured. Estimations were made by using energy balance analysis and the Penman–Monteith equation. Dew accumulation was also quantified using a cloth-plate method and by change in mass of whole plants. Tissue water content and total leaf area were similar in the two species, but maximum tank water content was three-fold greater for T. elongata than for T. brachycaulos, which had a lower degree of tissue desiccation tolerance than T. elongata. Succulence for T. brachycaulos changed little during most of the year, decreasing significantly only during one month in the dry season. Leaf water potential and osmotic pressure did not change significantly after a dew-deposition event. Theoretical dew per night on leaves was greatest during the early dry season (about 0.72 mm) compared with 0.34 mm in the rainy season and 0.008 mm in the late dry season. Dew amounts alone are not adequate to support growth but may maintain a favourable water balance for epiphytic bromeliads during the driest months of the year.

In tropical forests, severe droughts caused by El Niño events may strongly influence the water relations of tree seedlings and thereby increase their mortality. Data on known-aged seedlings of three common shade-tolerant canopy tree species (Trichilia tuberculata, Tetragastris panamensis and Quararibea asterolepis) in a Panamanian moist forest are presented. Seedling survival during a severe El Niño dry season (1997-98) was compared with prior long-term survival data, and levels of drought stress were assessed by measuring plant water potentials and gas exchange characteristics. Contrary to prediction, dry-season seedling survival was not dramatically reduced in any species compared with that expected in ‘normal’ years. In Trichilia and Quararibea, pre-dawn water potentials averaged −2 MPa and midday water potentials about −3 MPa. Stomatal conductances were very low, averaging 26 mmol m-2 s-1 for Tetragastris and 11–13 mmol m-2 s-1 for Trichilia and Quararibea. Photosynthetic rates also were very low but consistently positive, averaging 0.8–1.1 μmol m-2 s-1. The findings suggest that, once established, seedlings of common tree species in this semi-deciduous forest may be tolerant of drought events.

Summary 11

I. INTRODUCTION 12

II. LICHEN BIONT CHARACTERISTICS 12

1. The mycobiont 12

2. The photobiont 13

III. LICHEN GROWTH 13

1. Allocation of resources 13

2. Growth rates and environmental limitations 14

3. Maintaining an optimal energy use efficiency 15

IV. CARBON ACQUISITION 16

1. Water relations 16

(a) Desiccation tolerance 16

(b) Activation upon re-hydration 17

(c) Diffusion of water and CO2 18

2. Photobiont CO2 fixation 20

(a) CO2 acquisition modes 20

(b) Significance of the CCM 22

3. Light and nitrogen relations 22

(a) The light-response curve 23

(b) Photosynthetic capacity 25

(c) Coping with high light 27

V. CARBON SINKS AND EXPENDITURES 28

1. Carbon translocation 28

2. Carbon sinks 29

3. Respiration 30

VI. CONCLUDING REMARKS 30

Acknowledgements 31

References 31

Lichens are nutritionally specialized fungi (the mycobiont component) that derive carbon and in some cases nitrogen from algal or cyanobacterial photobionts. The mycobiont and photobiont live together in an integrated thallus, but they lack specific tissue for the transport of metabolites and resources between them. Carbon is acquired through photosynthesis in the photobiont, which is active when the lichen is wet and exposed to light. Lichen photosynthesis is limited primarily by water, light and nitrogen, but can also be constrained by slow diffusion of CO2 within the wet thallus. The assimilated carbon is exported from photobiont to mycobiont, which also predominates in terms of biomass, and apparently regulates the size of the photobiont population. It has therefore generally been assumed that most of the carbon is used for growth and maintenance of the fungal hyphae. However, the extent of photobiont respiration in relation to mycobiont respiration has seldom been quantified; neither do we know the pool sizes of various carbon sinks within lichens. Owing to this lack of fundamental data we do not know whether, or how, carbohydrate resources are regulated to maintain an optimal balance between energy input and expenditures in these symbiotic organisms. This review summarizes data on growth, carbon gain and carbon expenditures in lichens, with particular emphasis on factors determining the photosynthetic capacity of their photobionts. An attempt is made to introduce an economic analysis of lichen growth processes, a view that has often been adopted in studies of higher plants. Areas in which more data are needed for the construction of a model on ‘lichen resource allocation patterns’ are discussed.