Effects of the invasive plant Xanthium strumarium on diversity of native plant species: A competitive analysis approach in North and Northeast China.

Xanthium strumarium is native to North America and now has become one of the invasive alien species (IAS) in China. In order to detect the effects of the invader on biodiversity and evaluate its suitable habitats and ecological distribution, we investigated the abundance, relative abundance, diversity indices, and the number of the invasive and native plants in paired invaded and non-invaded quadrats in four locations in North and Northeast China. We also analyzed the effects of monthly mean maximum and minimum temperatures, relative humidity (%), and precipitations (mm). Strong positive significant (P < 0.01) correlation and maximum interspecific competition (41%) were found in Huailai between invaded and non-invaded quadrats. Shannon's Diversity Index showed that non-invaded plots had significantly (P < 0.05) more diversified species than invaded ones. The significant (P < 0.05) Margalef's Richness Index was found in Huailai and Zhangjiakou in non-invaded recorded heterogeneous nature of plant communities. Similarly, significant (P < 0.05) species richness found in Huailai and Zhangjiakou in non-invaded quadrats compared to invaded ones. Maximum evenness of Setaria feberi (0.47, 0.37), Seteria viridis (0.43) found in Fushun and Zhangjiakou recorded more stable in a community compared to other localities. Evenness showed positive relationship of Shannon Entropy within different plant species. The higher dissimilarity in plant communities found in Huailai (87.06%) followed by Yangyuan (44.43%), Zhangjiakou (40.13%) and Fushun (29.02%). The significant (P < 0.01) value of global statistics R (0.943/94.3%) showed high species diversity recorded in Huailai followed by Zhangjiakou recorded by non-metric multidimensional scaling and analysis of similarity between invaded and non-invaded plots. At the end it was concluded that the diversity indices reduced significantly (P < 0.05) in invaded quadrats indicated that native plant species become less diverse due to X. strumarium invasion. The degrees of X. strumarium invasion affected on species richness resulted to reduce diversity indices significantly in invaded quadrats.

Subcellular distribution and chemical forms involved in manganese accumulation and detoxification for Xanthium strumarium L.

Xanthium strumarium L. is a candidate species for manganese (Mn)-phyto-remediation. To reveal the mechanism of this species adaptive to Mn stress, the growth, Mn subcellular distribution, chemical forms, as well as micro-structure and ultra-structure responses of the mining ecotype (ME) of X. strumarium to Mn stress were studied with the non-mining ecotype (NME) as the reference by a hydroponic experiment. The results showed the ME demonstrated a higher tolerance to Mn stress with a superior growth and a higher tolerance index (TI) when compared with the NME. The concentrations of Mn in leaves, stems, and roots of the ME were 1.1-1.8, 1.2-1.9, and 1.3-1.9 times higher than those in the corresponding organs of the NME, respectively. The micro-structure and ultra-structure showed abnormal alterations, such as shrunken ducts and sieve canals, round-shaped chloroplasts, increased starch and osmiophilic granules, as well as expanded and non-compact granum thylakoids in the NME, compared to the ME. More than 83% of Mn was localized in cell wall and soluble fraction, while the Mn concentration in all fractions had a direct linear relationship with Mn treatment in the ME. The proportions of pectates and protein integrated-Mn, phosphate-Mn, and oxalate-Mn forms were dominant in leaves and stems of the ME, whereas, in the NME the relative proportions of inorganic Mn and water-soluble Mn forms in the roots was higher than the other forms. Altogether, the combination of preferential distribution of Mn in the cell wall and soluble fraction, and storage of Mn in low toxicity forms, such as phosphate-Mn, pectates and protein-bound Mn, and oxalate-Mn, might be responsible for alleviating Mn toxicity in the ME.

Effects of the mesophyll on stomatal responses in amphistomatous leaves.

The role of the mesophyll in stomatal functioning in thin amphistomatous leaves was investigated by altering gas exchange for one surface and observing the effects on stomatal conductance for the other surface. Three methods of perturbing gas exchange on the adaxial surface were used. First, gas exchange for the adaxial surface was completely blocked with plastic wrap or vacuum grease. Second, leaves were inverted to induce closure of the adaxial stomata. And third, ambient humidity for the adaxial surface was reduced to induce stomatal closure on that surface. Experiments were performed at low light intensity and three different CO2 concentrations to test the idea that stomatal responses in thin amphistomatous leaves are partially controlled by a signal from the mesophyll that varies with light and CO2 . In general, stomata on the abaxial surface opened when gas exchange on the adaxial surface was reduced, with the largest increases in conductance occurring at high CO2 concentration. The data are discussed with respect to role of a purported signal from the mesophyll and the partitioning of that signal between the two surfaces of the leaf.

The role of biomass allocation between lamina and petioles in a game of light competition in a dense stand of an annual plant.

Background and Aims: Models of plant three-dimensional (3-D) architecture have been used to find optimal morphological characteristics for light capture or carbon assimilation of a solitary plant. However, optimality theory is not necessarily useful to predict the advantageous strategy of an individual in dense stands, where light capture of an individual is influenced not only by its architecture but also by the architecture of its neighbours. Here, we analysed optimal and evolutionarily stable biomass allocation between the lamina and petiole (evolutionarily stable strategy; ESS) under various neighbour conditions using a 3-D simulation model based on the game theory. Methods: We obtained 3-D information of every leaf of actual Xanthium canadense plants grown in a dense stand using a ruler and a protractor. We calculated light capture and carbon assimilation of an individual plant when it stands alone and when it is surrounded by neighbours in the stand. We considered three trade-offs in petiole length and lamina area: biomass allocation, biomechanical constraints and photosynthesis. Optimal and evolutionarily stable biomass allocation between petiole and lamina were calculated under various neighbour conditions. Key Results: Optimal petiole length varied depending on the presence of neighbours and on the architecture of neighbours. The evolutionarily stable petiole length of plants in the stand tended to be longer than the optimal length of solitary plants. The mean of evolutionarily stable petiole length in the stand was similar to the real one. Trade-offs of biomechanical constraint and photosynthesis had minor effects on optimal and evolutionarily stable petiole length. Conclusion: Actual plants realize evolutionarily stable architecture in dense stands. Interestingly, there were multiple evolutionarily stable petiole lengths even in one stand, suggesting that plants with different architectures can coexist across plant communities.

Inconsistent intraspecific pattern in leaf life span along nitrogen-supply gradient.

PREMISE OF THE STUDY: Leaf life span (LLS) has long been hypothesized to plastically increase with decreasing nitrogen (N) supply from soil to maximize N retention, carbon assimilation, and fitness; however, accumulating evidence shows no consistent trend. The apparent inconsistencies are explained by a recent model that assumes LLS has a hump-shaped quadratic response to the N-supply gradient. The available evidence mostly originates from comparisons of LLS at only two levels of N availability, and the hypothesis remains unanswered. METHODS: We investigated LLS of two asteraceous forbs (Adenocaulon himalaicum and Xanthium canadense) experimentally grown at eight levels of N supply, which covered a range of N supply in their natural habitats. We additionally conducted a literature search to retrieve studies reporting LLS response along an N-supply gradient. KEY RESULTS: The LLS of neither species showed a hump-shaped response along the N-supply gradient. Past studies examining the LLS of an aquatic forb and terrestrial shrubs and trees along the N-supply gradient (more than four levels of N supply) also refuted the hypothesis. CONCLUSIONS: The LLS of a single species exhibited neither an increase nor a hump-shaped response to decreased N supply in a variety of life forms. Comparisons at only a few N levels are misleading with regard to LLS response to N supply.

Differential microRNA Analysis of Glandular Trichomes and Young Leaves in Xanthium strumarium L. Reveals Their Putative Roles in Regulating Terpenoid Biosynthesis.

The medicinal plant Xanthium strumarium L. (X. strumarium) is covered with glandular trichomes, which are the sites for synthesizing pharmacologically active terpenoids such as xanthatin. MicroRNAs (miRNAs) are a class of 21-24 nucleotide (nt) non-coding RNAs, most of which are identified as regulators of plant growth development. Identification of miRNAs involved in the biosynthesis of plant secondary metabolites remains limited. In this study, high-throughput Illumina sequencing, combined with target gene prediction, was performed to discover novel and conserved miRNAs with potential roles in regulating terpenoid biosynthesis in X. strumarium glandular trichomes. Two small RNA libraries from leaves and glandular trichomes of X. strumarium were established. In total, 1,185 conserved miRNAs and 37 novel miRNAs were identified, with 494 conserved miRNAs and 18 novel miRNAs being differentially expressed between the two tissue sources. Based on the X. strumarium transcriptome data that we recently constructed, 3,307 annotated mRNA transcripts were identified as putative targets of the differentially expressed miRNAs. KEGG (Kyoto Encyclopedia of Genes and Genomes) pathway analysis suggested that some of the differentially expressed miRNAs, including miR6435, miR5021 and miR1134, might be involved in terpenoid biosynthesis in the X. strumarium glandular trichomes. This study provides the first comprehensive analysis of miRNAs in X. strumarium, which forms the basis for further understanding of miRNA-based regulation on terpenoid biosynthesis.

Leaf dynamics in growth and reproduction of Xanthium canadense as influenced by stand density.

BACKGROUND AND AIMS: Leaf longevity is controlled by the light gradient in the canopy and also by the nitrogen (N) sink strength in the plant. Stand density may influence leaf dynamics through its effects on light gradient and on plant growth and reproduction. This study tests the hypothesis that the control by the light gradient is manifested more in the vegetative period, whereas the opposite is true when the plant becomes reproductive and develops a strong N sink. METHODS: Stands of Xanthium canadense were established at two densities. Emergence, growth and death of every leaf on the main stem and branches, and plant growth and N uptake were determined from germination to full senescence. Mean residence time and dry mass productivity were calculated per leaf number, leaf area, leaf mass and leaf N (collectively termed 'leaf variables') in order to analyse leaf dynamics and its effect on plant growth. KEY RESULTS: Branching and reproductive activities were higher at low than at high density. Overall there was no significant difference in mean residence time of leaf variables between the two stands. However, early leaf cohorts on the main stem had a longer retention time at low density, whereas later cohorts had a longer retention time at high density. Branch leaves emerged earlier and tended to live longer at low than at high density. Leaf efficiencies, defined as carbon export per unit investment of leaf variables, were higher at low density in all leaf variables except for leaf number. CONCLUSIONS: In the vegetative phase of plant growth, the light gradient strongly controls leaf longevity, whereas later the effects of branching and reproductive activities become stronger and over-rule the effect of light environment. As leaf N supports photosynthesis and also works as an N source for plant development, N use is pivotal in linking leaf dynamics with plant growth and reproduction.

Stem extension and mechanical stability of Xanthium canadense grown in an open or in a dense stand.

BACKGROUND AND AIMS: Plants in open, uncrowded habitats typically have relatively short stems with many branches, whereas plants in crowded habitats grow taller and more slender at the expense of mechanical stability. There seems to be a trade-off between height growth and mechanical stability, and this study addresses how stand density influences stem extension and consequently plant safety margins against mechanical failure. METHODS: Xanthium canadense plants were grown either solitarily (S-plants) or in a dense stand (D-plants) until flowering. Internode dimensions and mechanical properties were measured at the metamer level, and the critical buckling height beyond which the plant elastically buckles under its own weight and the maximum lateral wind force the plant can withstand were calculated. KEY RESULTS: Internodes were longer in D- than S-plants, but basal diameter did not differ significantly. Relative growth rates of internode length and diameter were negatively correlated to the volumetric solid fraction of the internode. Internode dry mass density was higher in S- than D-plants. Young's modulus of elasticity and the breaking stress were higher in lower metamers, and in D- than in S-plants. Within a stand, however, both moduli were positively related to dry mass density. The buckling safety factor, a ratio of critical buckling height to actual height, was higher in S- than in D-plants. D-plants were found to be approaching the limiting value 1. Lateral wind force resistance was higher in S- than in D-plants, and increased with growth in S-plants. CONCLUSIONS: Critical buckling height increased with height growth due mainly to an increase in stem stiffness and diameter and a reduction in crown/stem mass ratio. Lateral wind force resistance was enhanced due to increased tissue strength and diameter. The increase in tissue stiffness and strength with height growth plays a crucial role in maintaining a safety margin against mechanical failure in herbaceous species that lack the capacity for secondary growth.

Sesquiterpene lactone stereochemistry influences herbivore resistance and plant fitness in the field.

BACKGROUND AND AIMS: Stereochemical variation is widely known to influence the bioactivity of compounds in the context of pharmacology and pesticide science, but our understanding of its importance in mediating plant-herbivore interactions is limited, particularly in field settings. Similarly, sesquiterpene lactones are a broadly distributed class of putative defensive compounds, but little is known about their activities in the field. METHODS: Natural variation in sesquiterpene lactones of the common cocklebur, Xanthium strumarium (Asteraceae), was used in conjunction with a series of common garden experiments to examine relationships between stereochemical variation, herbivore damage and plant fitness. KEY RESULTS: The stereochemistry of sesquiterpene lactone ring junctions helped to explain variation in plant herbivore resistance. Plants producing cis-fused sesquiterpene lactones experienced significantly higher damage than plants producing trans-fused sesquiterpene lactones. Experiments manipulating herbivore damage above and below ambient levels found that herbivore damage was negatively correlated with plant fitness. This pattern translated into significant fitness differences between chemotypes under ambient levels of herbivore attack, but not when attack was experimentally reduced via pesticide. CONCLUSIONS: To our knowledge, this work represents only the second study to examine sesquiterpene lactones as defensive compounds in the field, the first to document herbivore-mediated natural selection on sesquiterpene lactone variation and the first to investigate the ecological significance of the stereochemistry of the lactone ring junction. The results indicate that subtle differences in stereochemistry may be a major determinant of the protective role of secondary metabolites and thus of plant fitness. As stereochemical variation is widespread in many groups of secondary metabolites, these findings suggest the possibility of dynamic evolutionary histories within the Asteraceae and other plant families showing extensive stereochemical variation.

Obsolete pesticides and application of colonizing plant species for remediation of contaminated soil in Kazakhstan.

In Kazakhstan, there is a problem of finding ways to clean local sites contaminated with pesticides. In particular, such sites are the deserted and destroyed storehouses where these pesticides were stored; existing storehouses do not fulfill sanitary standards. Phytoremediation is one potential method for reducing risk from these pesticides. Genetic heterogeneity of populations of wild and weedy species growing on pesticide-contaminated soil provides a source of plant species tolerant to these conditions. These plant species may be useful for phytoremediation applications. In 2008-2009 and 2011, we surveyed substances stored in 80 former pesticide storehouses in Kazakhstan (Almaty oblast) to demonstrate an inventory process needed to understand the obsolete pesticide problem throughout the country, and observed a total of 354.7 t of obsolete pesticides. At the sites, we have found organochlorine pesticides residues in soil including metabolites of dichlorodiphenyltrichloroethane and isomers of hexachlorocyclohexane. Twenty-four of the storehouse sites showed pesticides concentrations in soil higher than maximum allowable concentration which is equal to 100 µg kg(-1) in Kazakhstan. Seventeen pesticide-tolerant wild plant species were selected from colonizing plants that grew into/near the former storehouse's pesticides. The results have shown that colonizing plant annual and biannual species growing on soils polluted by pesticides possess ability to accumulate organochlorine pesticide residues and reduce pesticide concentrations in soil. Organochlorine pesticides taken up by the plants are distributed unevenly in different plant tissues. The main organ of organochlorine pesticide accumulation is the root system. The accumulation rate of organochlorine pesticides was found to be a specific characteristic of plant species and dependent on the degree of soil contamination. This information can be used for technology development of phytoremediation of pesticide-contaminated soils.

Mean residence time of leaf number, area, mass, and nitrogen in canopy photosynthesis.

Mean residence time (MRT) of plant nitrogen (N), which is an indicator of the expected length of time N newly taken up is retained before being lost, is an important component in plant nitrogen use. Here we extend the concept MRT to cover such variables as leaf number, leaf area, leaf dry mass, and nitrogen in the canopy. MRT was calculated from leaf duration (i.e., time integral of standing amount) divided by the total production of leaf variables. We determined MRT in a Xanthium canadense stand established with high or low N availability. The MRT of leaf number may imply longevity of leaves in the canopy. We found that the MRT of leaf area and dry mass were shorter than that of leaf number, while the MRT of leaf N was longer. The relatively longer MRT of leaf N was due to N resorption before leaf shedding. The MRT of all variables was longer at low N availability. Leaf productivity is the rate of canopy photosynthesis per unit amount of leaf variables, and multiplication of leaf productivity by MRT gives the leaf photosynthetic efficiency (canopy photosynthesis per unit production of leaf variables). The photosynthetic efficiency of leaf number implies the lifetime carbon gain of a leaf in the canopy. The analysis of plant-level N use efficiency by evaluating the N productivity and MRT is a well-established approach. Extension of these concepts to leaf number, area, mass, and N in the canopy will clarify the underlying logic in the study of leaf life span, leaf area development, and dry mass and N use in canopy photosynthesis.

Alternative perspective on the control of transpiration by radiation.

Stomatal responses to light are important determinants for plant water use efficiency and for general circulation models, but a mechanistic understanding of these responses remains elusive. A recent study [Pieruschka R, Huber G, Berry JA (2010) Proc Natl Acad Sci USA 107:13372-13377] concluded that stomata respond to total absorbed radiation rather than red and blue light as previously thought. We tested this idea by reexamining stomatal responses to red and blue light and to IR radiation. We show that responses to red and blue light are not consistent with a response to total absorbed radiation and that apparent stomatal responses to IR radiation are explainable as experimental artifacts. In addition, our data and analysis provide a method for accurately determining the internal temperature of a leaf.

Control of transpiration by radiation.

The terrestrial hydrological cycle is strongly influenced by transpiration--water loss through the stomatal pores of leaves. In this report we present studies showing that the energy content of radiation absorbed by the leaf influences stomatal control of transpiration. This observation is at odds with current concepts of how stomata sense and control transpiration, and we suggest an alternative model. Specifically, we argue that the steady-state water potential of the epidermis in the intact leaf is controlled by the difference between the radiation-controlled rate of water vapor production in the leaf interior and the rate of transpiration. Any difference between these two potentially large fluxes is made up by evaporation from (or condensation on) the epidermis, causing its water potential to pivot around this balance point. Previous work established that stomata in isolated epidermal strips respond by opening with increasing (and closing with decreasing) water potential. Thus, stomatal conductance and transpiration rate should increase when there is condensation on (and decrease when there is evaporation from) the epidermis, thus tending to maintain homeostasis of epidermal water potential. We use a model to show that such a mechanism would have control properties similar to those observed with leaves. This hypothesis provides a plausible explanation for the regulation of leaf and canopy transpiration by the radiation load and provides a unique framework for studies of the regulation of stomatal conductance by CO(2) and other factors.

Does leaf shedding increase the whole-plant carbon gain despite some nitrogen being lost with shedding?

When old leaves are shed, part of the nitrogen in the leaf is retranslocated to new leaves. This retranslocation will increase the whole-plant carbon gain when daily C gain : leaf N ratio (daily photosynthetic N-use efficiency, NUE) in the old leaf, expressed as a fraction of NUE in the new leaf, becomes lower than the fraction of leaf N that is resorbed before shedding (R(N)). We examined whether plants shed their leaves to increase the whole-plant C gain in accord with this criterion in a dense stand of an annual herb, Xanthium canadense, grown under high (HN) and low (LN) nitrogen availability. The NUE of a leaf at shedding expressed as a fraction of NUE in a new leaf was nearly equal to the R(N) in the LN stand, but significantly lower than the R(N) in the HN stand. Thus shedding of old leaves occurred as expected in the LN stand, whereas in the HN stand, shedding occurred later than expected. Sensitivity analyses showed that the decline in NUE of a leaf resulted primarily from a reduction in irradiance in the HN stand. On the other hand, it resulted from a reduction in irradiance and also in light-saturated photosynthesis : leaf N content ratio (potential photosynthetic NUE) in the LN stand.

How do leaf hydraulics limit stomatal conductance at high water vapour pressure deficits?

A reduction in leaf stomatal conductance (g) with increasing leaf-to-air difference in water vapour pressure (D) is nearly ubiquitous. Ecological comparisons of sensitivity have led to the hypothesis that the reduction in g with increasing D serves to maintain leaf water potentials above those that would cause loss of hydraulic conductance. A reduction in leaf water potential is commonly hypothesized to cause stomatal closure at high D. The importance of these particular hydraulic factors was tested by exposing Abutilon theophrasti, Glycine max, Gossypium hirsutum and Xanthium strumarium to D high enough to reduce g and then decreasing ambient carbon dioxide concentration ([CO2]), and observing the resulting changes in g, transpiration rate and leaf water potential, and their reversibility. Reducing the [CO2] at high D increased g and transpiration rate and lowered leaf water potential. The abnormally high transpiration rates did not result in reductions in hydraulic conductance. Results indicate that low water potential effects on g at high D could be overcome by low [CO2], and that even lower leaf water potentials did not cause a reduction in hydraulic conductance in these well-watered plants. Reduced g at high D in these species resulted primarily from increased stomatal sensitivity to [CO2] at high D, and this increased sensitivity may mediate stomatal responses to leaf hydraulics at high D.

Is evolution necessary for range expansion? Manipulating reproductive timing of a weedy annual transplanted beyond its range.

Ecologists often consider how environmental factors limit a species' geographic range. However, recent models suggest that geographic distribution also may be determined by a species' ability to adapt to novel environmental conditions. In this study, we empirically tested whether further evolution would be necessary for northern expansion of the weedy annual cocklebur (Xanthium strumarium) in its native North American range. We transplanted seedlings beyond the northern border and photoperiodically manipulated reproductive timing, a trait important for adaptation to shorter growing seasons at higher latitudes within the range, to determine whether further evolution of this trait would result in a phenotype viable beyond the range. Earlier reproductive induction enabled plants to produce mature seeds beyond the range and to achieve a reproductive output similar to those grown within the range. Therefore, evolution of earlier reproduction in marginal populations would be necessary for northward range expansion. This study is the first to empirically show that evolution in an ecologically important trait would enable a species to survive and reproduce beyond its current range. These results suggest that relatively few traits may limit a species' range and that identifying evolutionary constraints on such traits could be important for predicting geographic distribution.

Evidence for involvement of photosynthetic processes in the stomatal response to CO2.

Stomatal conductance (gs) typically declines in response to increasing intercellular CO2 concentration (ci). However, the mechanisms underlying this response are not fully understood. Recent work suggests that stomatal responses to ci and red light (RL) are linked to photosynthetic electron transport. We investigated the role of photosynthetic electron transport in the stomatal response to ci in intact leaves of cocklebur (Xanthium strumarium) plants by examining the responses of gs and net CO2 assimilation rate to ci in light and darkness, in the presence and absence of the photosystem II inhibitor 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU), and at 2% and 21% ambient oxygen. Our results indicate that (1) gs and assimilation rate decline concurrently and with similar spatial patterns in response to DCMU; (2) the response of gs to ci changes slope in concert with the transition from Rubisco- to electron transport-limited photosynthesis at various irradiances and oxygen concentrations; (3) the response of gs to ci is similar in darkness and in DCMU-treated leaves, whereas the response in light in non-DCMU-treated leaves is much larger and has a different shape; (4) the response of gs to ci is insensitive to oxygen in DCMU-treated leaves or in darkness; and (5) stomata respond normally to RL when ci is held constant, indicating the RL response does not require a reduction in ci by mesophyll photosynthesis. Together, these results suggest that part of the stomatal response to ci involves the balance between photosynthetic electron transport and carbon reduction either in the mesophyll or in guard cell chloroplasts.

Stress avoidance in a common annual: reproductive timing is important for local adaptation and geographic distribution.

Adaptation to local environments may be an important determinant of species' geographic range. However, little is known about which traits contribute to adaptation or whether their further evolution would facilitate range expansion. In this study, we assessed the adaptive value of stress avoidance traits in the common annual Cocklebur (Xanthium strumarium) by performing a reciprocal transplant across a broad latitudinal gradient extending to the species' northern border. Populations were locally adapted and stress avoidance traits accounted for most fitness differences between populations. At the northern border where growing seasons are cooler and shorter, native populations had evolved to reproduce earlier than native populations in the lower latitude gardens. This clinal pattern in reproductive timing corresponded to a shift in selection from favouring later to earlier reproduction. Thus, earlier reproduction is an important adaptation to northern latitudes and constraint on the further evolution of this trait in marginal populations could potentially limit distribution.

Atmospheric CO2 enrichment alters energy assimilation, investment and allocation in Xanthium strumarium.

Energy-use efficiency and energy assimilation, investment and allocation patterns are likely to influence plant growth responses to increasing atmospheric CO2 concentration ([CO2]). Here, we describe the influence of elevated [CO2] on energetic properties as a mechanism of growth responses in Xanthium strumarium. Individuals of X. strumarium were grown at ambient or elevated [CO2] and harvested. Total biomass and energetic construction costs (CC) of leaves, stems, roots and fruits and percentage of total biomass and energy allocated to these components were determined. Photosynthetic energy-use efficiency (PEUE) was calculated as the ratio of total energy gained via photosynthetic activity (Atotal) to leaf CC. Elevated [CO2] increased leaf Atotal, but decreased CC per unit mass of leaves and roots. Consequently, X. strumarium individuals produced more leaf and root biomass at elevated [CO2] without increasing total energy investment in these structures (CCtotal). Whole-plant biomass was associated positively with PEUE. Whole-plant construction required 16.1% less energy than modeled whole-plant energy investment had CC not responded to increased [CO2]. As a physiological mechanism affecting growth, altered energetic properties could positively influence productivity of X. strumarium, and potentially other species, at elevated [CO2].

MSMA resistance studies.

Monosodium methanearsonate (MSMA)-resistant and -susceptible common cocklebur (Xanthium strumarium L.) and cotton (Gossypium hirsutum L.) were treated with MSMA. Plant parameters analyzed were: glutathione synthetase activity, selected amino acid (arginine, glutamic acid, alanine, citrulline, glutamine, and glutathione) content and arsenic content (MSMA, total arsenic, and arsonate). No reduction of arsenic from the parent pentavalent form present in MSMA to the trivalent form was detected. Arginine, glutamic acid, and glutamine content increased in tissue three days after MSMA treatment. Glutathione content decreased during the first three days after treatment; however, five days after treatment the resistant biotype of cocklebur and cotton had elevated glutathione levels (8-20 times greater, respectively). Glutathione Synthetase activity was higher in cotton than in either of the cocklebur biotypes; MSMA did not affect its activity in cotton or either cocklebur biotype. Resistant biotypes have a slightly higher activity than the susceptible biotype. Tolerance of cotton to MSMA may be related to glutathione synthetase activity and possibly to the presence of phytochelatins. Also, increased glutathione levels in the resistant biotype may implicate phytochelatin involvement in the resistance mechanism.