Effects of litter amount and seed sowing position on seedling emergence and growth of hemiparasitic Rhinanthus species under drought stress.

Roadside vegetation in Central Europe is mostly species-poor and dominated by a few grass species. Hemiparasitic plant species, including Rhinanthus spp., might effectively restrict grass growth, thereby making space for light-dependent herb species. Despite the significance of abiotic site conditions for plant establishment in general, their effects on Rhinanthus establishment are less well known. We investigated combined effects of water availability, litter amount and seed position within litter on Rhinanthus seedling emergence and growth. Two parallel greenhouse experiments were conducted with R. angustifolius and R. minor. In these, we tested the impact of 200 or 400 g litter·m-2 with seeds sown beneath or on top of a litter layer under constantly humid or intermittently dry conditions on seedling emergence and biomass production of Rhinanthus. Presence of litter positively affected Rhinanthus seedling emergence when sown beneath the litter layer and reduced negative effects of water deficiency. Sowing beneath a litter layer increased seedling emergence by 157%, with similar effects at 200 and 400 g litter·m-2. Water level did not affect biomass production. Compared to R. minor, R. angustifolius had higher mean biomass, and its seedlings emerged earlier and in higher numbers. Our results indicate that Rhinanthus spp. react similarly to litter as non-hemiparasitic plant species from temperate grasslands. Litter presence positively influenced Rhinanthus seedling emergence and growth under intermittently dry conditions. Its hemiparasitic characteristics might reduce drought impacts on biomass production. To ensure seed contact with the soil surface, seeds should be sown when no litter is present, or mulching should occur post-sowing.

Interannual variation in season length is linked to strong co-gradient plasticity of phenology in a montane annual plant.

Species are commonly distributed along latitudinal and elevational gradients of growing season length to which they might respond via phenotypic plasticity and/or adaptive genetic differentiation. However, the relative contribution of these processes and whether plasticity, if it occurs, facilitates expansion along season-length gradients remain unclear, but are important for predicting species fates during anthropogenic change. We quantified phenological trait variation in the montane annual Rhinanthus minor for three generations at 12 sites across 900 m of elevation in the Canadian Rocky Mountains and conducted a reciprocal transplant experiment for two generations among nine sites. We compared clines and interannual variation of phenological traits between natural and transplanted individuals. Season length declined by c. 37% along our elevational gradient and, as expected, plants emerged, reached first flower and made their first seed in c. 41% fewer growing degree days under shorter growing seasons. Although reciprocal transplants revealed modest genetic differentiation across elevation, trait clines primarily were due to striking co-gradient plasticity that paralleled genetic differentiation. Co-gradient plasticity likely evolved in response to considerable interannual variation in season length across our elevational transect, and should prepare R. minor to make adaptive changes to phenology in response to ongoing climate change predicted for montane environments.

Local adaptation primes cold-edge populations for range expansion but not warming-induced range shifts.

According to theory, edge populations may be poised to expand species' ranges if they are locally adapted to extreme conditions, or ill-suited to colonise beyond-range habitat if their offspring are genetically and competitively inferior. We tested these contrasting predictions by transplanting low-, mid-, and high-elevation (edge) populations of an annual plant throughout and above its elevational distribution. Seed from poor-quality edge habitat (one of two transects) had inferior emergence, but edge seeds also had adaptive phenology (both transects). High-elevation plants flowered earlier, required less heat accumulation to mature seed, and so achieved higher lifetime fitness at and above the range edge. Experimental warming improved fitness above the range, but eliminated the advantage of local cold-edge populations, supporting recent models in which cold-adapted edge populations do not facilitate warming-induced range shifts. The highest above-range fitness was achieved by a 'super edge phenotype' from a neighbouring mountain, suggesting key adaptations exist regionally even if absent from local edge populations.

Arabinogalactan protein-rich cell walls, paramural deposits and ergastic globules define the hyaline bodies of rhinanthoid Orobanchaceae haustoria.

BACKGROUND AND AIMS: Parasitic plants obtain nutrients from their hosts through organs called haustoria. The hyaline body is a specialized parenchymatous tissue occupying the central parts of haustoria in many Orobanchaceae species. The structure and functions of hyaline bodies are poorly understood despite their apparent necessity for the proper functioning of haustoria. Reported here is a cell wall-focused immunohistochemical study of the hyaline bodies of three species from the ecologically important clade of rhinanthoid Orobanchaceae. METHODS: Haustoria collected from laboratory-grown and field-collected plants of Rhinanthus minor, Odontites vernus and Melampyrum pratense attached to various hosts were immunolabelled for cell wall matrix glycans and glycoproteins using specific monoclonal antibodies (mAbs). KEY RESULTS: Hyaline body cell wall architecture differed from that of the surrounding parenchyma in all species investigated. Enrichment in arabinogalactan protein (AGP) epitopes labelled with mAbs LM2, JIM8, JIM13, JIM14 and CCRC-M7 was prominent and coincided with reduced labelling of de-esterified homogalacturonan with mAbs JIM5, LM18 and LM19. Furthermore, paramural bodies, intercellular deposits and globular ergastic bodies composed of pectins, xyloglucans, extensins and AGPs were common. In Rhinanthus they were particularly abundant in pairings with legume hosts. Hyaline body cells were not in direct contact with haustorial xylem, which was surrounded by a single layer of paratracheal parenchyma with thickened cell walls abutting the xylem. CONCLUSIONS: The distinctive anatomy and cell wall architecture indicate hyaline body specialization. Altered proportions of AGPs and pectins may affect the mechanical properties of hyaline body cell walls. This and the association with a transfer-like type of paratracheal parenchyma suggest a role in nutrient translocation. Organelle-rich protoplasts and the presence of exceptionally profuse intra- and intercellular wall materials when attached to a nitrogen-fixing host suggest subsequent processing and transient storage of nutrients. AGPs might therefore be implicated in nutrient transfer and metabolism in haustoria.

The genetics of indirect ecological effects-plant parasites and aphid herbivores.

When parasitic plants and aphid herbivores share a host, both direct and indirect ecological effects (IEEs) can influence evolutionary processes. We used a hemiparasitic plant (Rhinanthus minor), a grass host (Hordeum vulgare) and a cereal aphid (Sitobion avenae) to investigate the genetics of IEEs between the aphid and the parasitic plant, and looked to see how these might affect or be influenced by the genetic diversity of the host plants. Survival of R. minor depended on the parasite's population of origin, the genotypes of the aphids sharing the host and the genetic diversity in the host plant community. Hence the indirect effects of the aphids on the parasitic plants depended on the genetic environment of the system. Here, we show that genetic variation can be important in determining the outcome of IEEs. Therefore, IEEs have the potential to influence evolutionary processes and the continuity of species interactions over time.

The impact of phosphorus on interactions of the hemiparasitic angiosperm Rhinanthus minor and its host Lolium perenne.

The effects of phosphorus supply on the outcome of interactions between the hemiparasitic angiosperm Rhinanthus minor L. with its host species Lolium perenne L. were investigated in a glasshouse experiment. Host plants were grown in 3-l pots in the presence and absence of R. minor at limiting (0.13 mM P) and optimal (0.65 mM P) concentrations of phosphorus for the growth of the host species. Phosphorus was supplied at 2-day intervals in the form of half-strength Long Ashton nitrate-based solution with phosphorus concentrations adjusted accordingly. Parasitism by R. minor significantly suppressed host growth, with final biomass losses ranging between 32% and 44%. Phosphorus supply had a marked impact on the outcome of the host-parasite interaction. By the end of the growing period, parasite biomass at 0.65 mM P was 90% lower than that achieved at 0.13 mM P. In contrast, host biomass at 0.65 mM P was 74% higher than achieved at 0.13 mM P, indicting that the negative impact of parasitism on the host species was reduced when phosphorus supply was increased. The effects of phosphorus on the host-parasite association appeared to be mediated by changes in both the morphological characteristics of the host roots and the relative sink strengths of the host and parasite.