Climate, phylogeny and the ecological distribution of C4 grasses

'C4 photosynthesis' refers to a suite of traits that increase photosynthesis in high light and high temperature environments. Most C4 plants are grasses, which dominate tropical and subtropical grasslands and savannas but are conspicuously absent from cold growing season climates. Physiological attributes of C4 photosynthesis have been invoked to explain C4 grass biogeography; however, the pathway evolved exclusively in grass lineages of tropical origin, suggesting that the prevalence of C4 grasses in warm climates could be due to other traits inherited from their non-C4 ancestors. Here we investigate the relative influences of phylogeny and photosynthetic pathway in determining the ecological distributions of C4 grasses in Hawaii. We find that the restriction of C4 grasses to warmer areas is due largely to their evolutionary history as members of a warm-climate grass clade, but that the pathway does appear to confer a competitive advantage to grasses in more arid environments.     

Consider the grasses

The identification of grasses relies heavily upon the structure ofspikelets, which, at first sight, appear to be very complicated. This article attempts to show that this complexity is more apparent than real. Spikelets display an enormous range in the shape and peculiar modifications of their separate parts, but all are constructed on the same basic plan. By relating the unfamiliar terminology for spikelet parts with that for more conventional inflorescences, and by showing how spikelets which look very different can be interpreted as variations on one basic theme, it is hoped that the initial difficulties experienced by many people when they attempt to name grasses will be largely overcome.     

Correlated rates of synonymous site evolution across plant genomes

Synonymous substitution rates have been shown to vary among evolutionary lineages of both nuclear and organellar genes across a broad range of taxonomic groups. In animals, rate heterogeneity does not appear to be correlated across nuclear and mitochondrial genes. In this paper, we contrast substitution rates in two plant groups and show that grasses evolve more rapidly than palms at synonymous sites in a mitochondrial, a nuclear, and a plastid gene. Furthermore, we show that the relative rates of synonymous substitution between grasses and palms are similar at the three loci. The correlation in synonymous substitution rates across genes is particularly striking because the three genes evolve at very different absolute rates. In contrast, relative rates of nonsynonymous substitution are not conserved among the three genes.      

ENDOPHYTE-HOST ASSOCIATIONS IN GRASSES. XVI. PATTERNS OF ENDOPHYTE DISTRIBUTION IN SPECIES OF THE TRIBE AGROSTIDEAE

Distribution of Acremonium endophytes in several species of Agrostideae is determined through surveys conducted in the field and in grass herbaria. Widespread geographic distribution of Acremonium in a particular grass species may indicate early colonization of the grass as a host, while a narrow pattern of geographic distribution suggests more recent colonization of the grass. Two grasses that appear to show a narrow endophyte distribution include Agrostis alba and Ammophila breviligulata. Cultural features of endophytes from several grasses are compared. A new variety of Acremonium typhinum is proposed to accommodate the endophyte from A. breviligulata. An endophyte in Agrostis alba is identified as Acremonium starrii. It is also suggested that Acremonium endophytes are spreading in grasses and may be progressively colonizing new hosts.     

A rodent plague on prairie diversity

Selective vole ( Microtus pennsylvanicus ) suppression of prairie grasses and forbs in experimental restorations suggests why many of the plants are likely to be uncommon in nature. Vole herbivory reduced densities of legumes and grasses and increased unpalatable forbs in replicated plantings in Illinois: six otherwise common species ( Dalea purpurea , Desmanthus illinoensis , Elymus canadensis , Panicum virgatum , Phalaris arundinacea , Sorghastrum nutans ) declined 27–89% in abundance, whereas two species ( Echinacea purpurea and Rudbeckia hirta ) increased by 61% and 1023%. Species number dropped by 19% and plant diversity (Simpson's D) by 37% in one treatment to which voles had access. Plots were planted with 18 prairie species of the region, but in even distributions of 35 or 350 seeds species⁻¹ m⁻², rather than skewed in favour of large C₄ grasses common in native remnants. Manipulation of plant composition and vole access revealed what are likely to be formative effects of rodent herbivory on vegetative composition. These experimental tallgrass communities appear to be assembling from plant species that voles prefer not to eat.     

Non-indigenous grasses impede woody succession

With the proliferation of old fields and the decline of native grasslands in North America, non-indigenous grasses, which tend to colonize and dominate North American old fields, have become progressively more abundant. These new grasses can differ from native grasses in a number of ways, including root and shoot morphology (e.g., density of root mat, height of shoots), growth phenology (e.g., cool season vs. warm season growth), and plant–soil–water relations due to differences in photosynthetic physiology (C₃ vs. C₄). Woody plants have been slow to colonize some old fields in the prairie-forest border area of North America and it is hypothesized that non-indigenous grasses may be contributing to the poor establishment success of woody plants in this region, possibly through more intense competition for resources. To test this hypothesis, a multi-factorial field experiment was conducted in which water, nitrogen, and grass functional group (non-indigenous C₃ and native C₄ species) were manipulated in a study of survival of oak seedlings. The grass type variously affected some of the different growth measurements, however, the effects of grass type on seedling growth were small compared to the effects on seedling survival. The results showed that when grown under dry conditions, seedlings growing in non-indigenous grasses experienced up to a 50% reduction in survival compared to those growing in native grasses under the same conditions. Analyses of root and shoot competition showed that the cause for the reduced survival in the non-indigenous grasses was due primarily to underground processes. The findings confirmed our initial hypothesis that non-indigenous grasses are likely contributing to the poor establishment success of woody plants in these old fields. However, the explanation for the reduced oak seedling survival in non-indigenous grasses does not appear to be due to reduced resource availability since soil water levels did not differ between non-indigenous and native grass plots and other resource levels measured (light, NO₃, and NH₄) were higher in non-indigenous grass plots under dry conditions. An alternative explanation is that the non-indigenous grasses modify the soil environment in ways that, under dry conditions, are deleterious to emerging oak seedlings. Since current climate projections for the upper Midwest are for hotter and drier summers, the results suggest that the resistance of these old fields to oak encroachment will likely increase in the future.     

Complex interactions between a legume and two grasses in a subalpine meadow

Interactions between plants are a complex combination of positive and negative interactions, with the net outcome depending on environmental contexts. The more frequent association of Trifolium alpinum (legume) with Festuca eskia than with Nardus stricta (grasses) in many Pyrenean subalpine meadows suggests a differential ability to use nitrogen (N) derived from N(2) fixation. In the field, we investigated the interactions between the legume and grasses and, in the glasshouse, the transfer of (15)N from the legume to the grasses. In one grass-Trifolium mixture, the legume had a strong positive effect on the biomass and N content of the grass as compared to pure grass stands. When both grasses grew together with the legume, only Festuca benefited from the presence of Trifolium but, surprisingly, the benefit decreased with increasing Trifolium abundance. Leaf labeling experiments with (15)N-NH(4)(+) revealed a higher transfer of (15)N from Trifolium to Festuca than to Nardus, suggesting a more direct N pathway between the two species. This more direct pathway could prevent Nardus from benefiting from the legume N in the three-species mixtures. Thus, the positive interactions between N-fixers and nonfixers appear to be largely species-specific and to depend strongly on the species in the plant assemblage.     

Grasses as appropriate targets in weed biocontrol: is the common reed, <Emphasis Type="Italic">Phragmites australis</Emphasis>, an anomaly?

Despite their importance as invasive species, there has been a hesitation to target grasses in classical biocontrol. This historic bias appears to be changing with multiple active research and release programs. Similarly, biocontrol workers appear to avoid targeting species with native congeners. These biases appear inappropriate as the ecological and entomological literature provide abundant evidence for sub-genus specificity for many herbivores, including those attacking grasses. The biocontrol program targeting Phragmites australis (Cav.) Trin. ex Steud (Poaceae) provides an informative example with endemic subspecies in North America and many sub-genus specific herbivores, including potential European control agents. Grasses and target weeds with congeneric native species require rigorous host range testing, similar to all other targets in current weed biological control programs. Furthermore, it appears prudent to ask petition reviewers and regulatory agencies to abandon their focus on results of no-choice studies and to distinguish between trivial feeding and demographic impacts.     

Strong population structure characterizes weediness gene evolution in the invasive grass species Brachypodium distachyon

Mediterranean annual grasses have invaded California and have replaced vast areas of native grassland. One of these invasive grasses is Brachypodium distachyon , a new model species for the grasses with extensive genomic resources and a nearly completed genome sequence. This study shows that the level of genetic variation in invaded California grasslands is lower compared to the native range in Eurasia. The invaded regions are characterized by highly differentiated populations of B. distachyon isolated by distance, most likely as a result of founder effects and a dearth of outcrossing events. EXP6 and EXP10 encoding α-expansins responsible for rapid growth, and AGL11 and AGL13 encoding proteins involved in vegetative phase regulation, appear to be under purifying selection with no evidence for local adaptation. Our data show that B. distachyon has diverged only recently from related Brachypodium species and that tetraploidization might have been as recent as a few thousand years ago. Observed low genetic variation in EXP10 and AGL13 appears to have been present in Eurasia before tetraploidization, potentially as a result of strong selective pressures on advantageous mutations, which are most likely responsible for its fast growth and rapid completion of its life cycle.     

Mind the gap among patches in arid plant communities: rapid root proliferation in response to N addition

Aims In arid communities, it has been proposed that individual plants can extend their roots beyond their canopy exploring neighbouring bare ground areas. This becomes relevant in systems where the vegetation is distributed in patches surrounded by bare soil. However, whether roots of different species may be overlapping under bare ground areas is still controversial. The factors controlling root responses when no plants appear to be directly influencing the gap among patches are still unclear. The aim of our study was to detect perennial grasses responses to an N enrichment pulse.     

C4 grasses adapted to low precipitation habitats show traits related to greater mesophyll conductance and lower leaf hydraulic conductance

In habitats with low water availability, a fundamental challenge for plants will be to maximize photosynthetic C-gain while minimizing transpirational water-loss. This trade-off between C-gain and water-loss can in part be achieved through the coordination of leaf-level photosynthetic and hydraulic traits. To test the relationship of photosynthetic C-gain and transpirational water-loss, we grew, under common growth conditions, 18 C₄ grasses adapted to habitats with different mean annual precipitation (MAP) and measured leaf-level structural and anatomical traits associated with mesophyll conductance (g_m) and leaf hydraulic conductance (K_leaf). The C₄ grasses adapted to lower MAP showed greater mesophyll surface area exposed to intercellular air spaces (S_mes) and adaxial stomatal density (SD_ada) which supported greater g_m. These grasses also showed greater leaf thickness and vein-to-epidermis distance, which may lead to lower K_leaf. Additionally, grasses with greater g_m and lower K_leaf also showed greater photosynthetic rates (A_net) and leaf-level water-use efficiency (WUE). In summary, we identify a suite of leaf-level traits that appear important for adaptation of C₄ grasses to habitats with low MAP and may be useful to identify C₄ species showing greater A_net and WUE in drier conditions.     

Mycorrhizal suppression alters plant productivity and forb establishment in a grass-dominated prairie restoration

A fundamental goal of restoration is the re-establishment of plant diversity representative of native vegetation. However, many prairie restorations or Conservation Reserve Program sites have been seeded with warm-season grasses, leading to grass-dominated, low-diversity restorations not representative of native grasslands. These dominant grasses are strongly mycotrophic, while many subordinate forb species appear to be less dependent on mycorrhizal symbiosis. Therefore, manipulating arbuscular mycorrhizal fungi (AMF) may be useful in promoting establishment and growth of forb species in grass-dominated prairie restorations. To assess the potential role of mycorrhizae in affecting the productivity and community composition of restored tallgrass prairie, we conducted a 4-year field experiment on an 8-year-old grassland restoration at the Konza Prairie in northeastern Kansas, USA. At the initiation of our study, seeds of 12 forb species varying in degree of mycorrhizal dependence were added to established grass-dominated plots. Replicate plots were treated bi-weekly with a soil drench of fungicide (Topsin-M^®) over four growing seasons and compared to non-treated control plots to assess the role of AMF in affecting plant species composition, productivity, leaf tissue quality, and diversity in restored tallgrass prairie. Topsin applications successfully reduced mycorrhizal colonization of grass roots to approximately 60–80% relative to roots in control plots. Four years of mycorrhizal suppression reduced productivity of the dominant grasses and increased plant species richness and diversity. These results highlight the importance of mycorrhizae as mediators of plant productivity and community dynamics in restored tallgrass prairie and indicate that temporarily suppressing AMF decreases productivity of the dominant C4 grasses and allows for establishment of seeded forb species.     

Flowering and determinacy in maize

All plant organs are produced by meristems, groups of stem cells located in the tips of roots and shoots. Indeterminate meristems make an indefinite number of organs, whereas determinate meristems are consumed after making a specific number of organs. Maize is an ideal system to study the genetic control of meristem fate because of the contribution from determinate and indeterminate meristems to the overall inflorescence. Here, the latest work on meristem maintenance and organ specification in maize is reviewed. Genetic networks, such as the CLAVATA components of meristem maintenance and the ABC programme of flower development, are conserved between grasses and eudicots. Maize and rice appear to have conserved mechanisms of meristem maintenance and organ identity. Other pathways, such as sex determination, are likely to be found only in maize with its separate male and female flowers. A rich genetic history has resulted in a large collection of maize mutants. The advent of genomic tools and synteny across the grasses now permits the isolation of the genes behind inflorescence architecture and the ability to compare function across the Angiosperms.     

Systematics, distribution, and host specificity of grass endophytes.

Clavicipitaceous endophytes (Ascomycetes) are distributed worldwide in many grasses and sedges forming a perennial and often mutualistic association with their hosts. Most endophytes appear to produce alkaloid toxins in infected plants. The high frequency of infection in many grasses and in certain grassland communities may indicate a selective advantage of infected over non-infected host plants due to their toxic effects on grazing animals and insects. Field observations and artificial inoculations of seedlings have demonstrated a high degree of specificity of most endophytes to their host plant, particularly in asexual, seed-borne endophytes. Specific isozyme genotypes found on several host species suggest that host-specific physiological races may occur. Knowledge of host range and host specificity is vital for potential applications of endophytes in pest control.     

Temporal and micro-spatial patterning of seedling establishment. Consequences for patch dynamics in the southern Monte,Argentina

We compared the temporal and micro-spatial patterning of seedling emergence and establishment of two cohorts of perennial grasses and shrubs in the southern Monte, Argentina. Samplings were carried out in two contrasting areas (grazed and non-grazed) during four years. We found lower densities of emerged and established seedlings of perennial grasses in the grazed relative to the non-grazed area. Conversely, emerged seedlings of shrubs did not vary between the grazed and the non-grazed area and densities of established shrub seedlings were higher in the grazed than in the non-grazed area. We only found differences between cohorts in seedling emergence of perennial grasses. These differences could be associated with the amount of precipitation in the year previous to the emergence. Both in the grazed and non-grazed area, seedlings of perennial grasses were concentrated at the periphery of plant patches. We defined a plant patch as a discrete unit of the spatial pattern of vegetation surrounded by, at least, 20 cm of bare soil from the nearest neighbour patch. Emergence in perennial grasses was more frequent at the southern/western patch-periphery than at other patch-periphery locations. Established seedlings of perennial grasses, however, were homogeneously distributed throughout patch periphery. Emergence in shrubs was more frequent at the centre and periphery of patches than at inter-patch microsites. In contrast, established seedlings of shrubs were homogeneously distributed among microsites. Our results suggests that differential seedling establishment between life forms is the outcome of complex biotic and abiotic interactions and feedbacks at the patch level between seedlings and established plants. Both life forms appear to have a different role in the origin, dynamics, and maintenance of spotting vegetation. Because of the ability to establish both at inter-patch and patch microsites, shrubs could be identified as colonizers or initiators of small plant patches in bare soil or they may contribute to increase the cover and size of pre-existing plant patches. Both processes would be promoted in grazed areas. Due to the ability to establish at patch peripheries, perennial grasses would contribute to the isodiametric growth of pre-existing patches, preferentially in non-grazed areas. This revised version was published online in June 2006 with corrections to the Cover Date.     

Seasonal and species variation in the content of cadmium and associated metals in pasture plants at Shipham

Summary Very high Cd and Zn concentrations exist in soils reclaimed from old mine workings at Shipham but little of the metal is transferred into the pasture herbage. The amount of metal in the soil therefore only influences the amount in the aerial part of the plant to a small degree. It would appear to be the plant species which to a large extent governs its metal burden: grasses accumulating the least Cd and members of the Compositae the most.Within this species variation exists a change in metal content that corresponds to the time of year, setal levels in plants peaking in the winter between January and March. This increase in the metal content of the shoot reflects the redistribution of metal previously bound within the root. The movement of Pb may be associated with changes in the phosphate status of the plant. In grasses it would appear that the root possesses the highest metal burden, and forHolcus lanatus, tolerance to both Cd and Zn has been established. The ammonium-acetate and DTPA-extractable Zn/Cd ratio in soils corresponds closely to that found in the roots of both ryegrass and Yorkshire fog. A much higher ratio was observed in the shoots of these plants.     

The chloroplast generps 4 as a tool for the study ofPoaceae phylogeny

Phylogenetic analyses of 28Poaceae species based on the chloroplastrps 4 gene are presented using parsimony and distance methods. Two monocots from other families were used as outgroups. The chloroplast generps 4 was amplified, cloned, and sequenced for each species. The inferred phylogenetic trees were compared to recent classifications and are shown to fit their general features. There is a dichotomy in our tree between the pooid group and the other grasses. This is in contradiction with other molecular phylogenies, where the bamboos appear first within the family. This result led us to discuss some hypotheses about the relationships of the bambusoids with the other groups of grasses, and also about the relative position of rice and bamboo, which are found close to each other in our trees.     

Abundance and diaspore weight in rare and common prairie grasses

Summary Abundance (g/m²) and diaspore weight are positively correlated in seven species of perennial grasses that occur in prairies. The rare grasses (<10.0 g/m²) have light dispersal units (0.06 to 1.76 mg); the common grasses (>10.0 g/m²) have heavy dispersal units (2.23 to 2.80 mg). This result is the first reported differentiating trait between related rare and common organisms occurring in same habitat.Three hypotheses that explain this phenomenon are compared; the third most likely holds. First, rare grasses may be rare because their small seeds are less successful in establishment than those of common grasses. Second, if the persistence of small populations is marginal, rare grasses may devote less energy (or other currency) to seed production. Third, rare grasses may be colonizers of spatially and temporally rare microsites appropriate for growth and thus have seeds adapted for longer distance dispersal than those of common grasses. This last hypothesis suggests a new pathway for the evolution of weeds.