Shoot: root ratio of seedlings is associated with species niche on soil moisture gradient.

Surviving the seedling phase is crucial for the establishment of plant individuals and populations. In ecosystems with dynamic water availability such as temperate grasslands, seedlings should adjust their growth strategy not only to match the current conditions but also to secure resource acquisition in the future. Here, we explored evolutionary adaptations determining plant early growth strategies in herbaceous species of temperate grasslands differing in their requirements for soil water availability. We chose 15 plant genera, within which we selected species differing in their Ellenberg indicator values for moisture. We cultivated the seedlings under standard conditions with sufficient water supply for 4 weeks. Subsequently, we measured length-based and mass-based shoot:root ratio to investigate seedling growth strategy and its association with species ecological niche. Seed size and content of soil-borne nutrients were considered as potential covariates affecting this association. Linear mixed-effect models identified the length-based shoot:root ratio of seedlings was positively associated with soil moisture requirements in a congeneric species comparison. Nitrogen and phosphorus seed concentrations had an additional negative effect on the shoot:root ratio. Neither of these trends was found for the mass-based shoot:root ratio. We demonstrated for the first time that there might be a general adaptation modifying the seedling shoot:root ratio according to the species niche position on the soil moisture gradient in temperate grassland species across a broad range of angiosperm phylogeny. This adaptation seems to be affected by seed mineral nutrient reserves and may operate in parallel to the well-known phenotypic plasticity.

Thesium linophyllon parasitizes and suppresses expansive Calamagrostis epigejos.

Root-hemiparasitic interaction between the dominant grass Calamagrostis epigejos and the hemiparasite Thesium linophyllon was studied to assess the potential of the parasite to regulate dominance of the grass, which is expanding into species-rich steppe grasslands. First, we aimed to identify physiological links between the two species as a principal indicator of the parasitic relationship. Second, we analysed the dynamics of the two species in the vegetation of a steppe grassland at the foot of the Bükk Mountains, Hungary, where their joint presence is recorded in a long-term permanent plot monitoring dataset to detect patterns associated with the parasitic ecological interaction. Numerous well-developed functional haustoria of Th. linophyllon were identified on the root systems of C. epigejos. The joint dynamics of C. epigejos and Th. linophyllon displayed clear signs of the parasitic interaction: (1) the dynamics of Th. linophyllon frequency was positively associated with the initial cover of C. epigejos; (2) maximum recorded cover values of the two species were strongly positively correlated; and (3) the extent of C. epigejos decrease in the vegetation was significantly positively associated with maximum Th. linophyllon cover recorded throughout the monitoring period. We demonstrate that C. epigejos can be parasitized by Th. linophyllon, which restricts abundance of the grass. Th. linophyllon thus has potential to act as a native biological control of C. epigejos in steppe grasslands.

A hemiparasite in the forest understorey: photosynthetic performance and carbon balance of Melampyrum pratense.

Melampyrum pratense is an annual root-hemiparasitic plant growing mostly in forest understorey, an environment with unstable light conditions. While photosynthetic responses of autotrophic plants to variable light conditions are in general well understood, light responses of root hemiparasites have not been investigated. We carried out gas exchange measurements (light response and photosynthetic induction curves) to assess the photosynthetic performance of M. pratense in spring and summer. These data and recorded light dynamics data were subsequently used to model carbon balance of the hemiparasite throughout the entire growth season. Summer leaves had significantly lower rates of saturated photosynthesis and dark respiration than spring leaves, a pattern expected to reflect the difference between sun- and shade-adapted leaves. However, even the summer leaves of the hemiparasite exhibited a higher rate of light-saturated photosynthesis than reported in non-parasitic understorey herbs. This is likely related to its annual life history, rare among other understorey herbs. The carbon balance model considering photosynthetic induction still indicated insufficient autotrophic carbon gain for seed production in the summer months due to limited light availability and substantial carbon loss through dark respiration. The results point to potentially high importance of heterotrophic carbon acquisition in M. pratense, which could be of at least comparable importance as in other mixotrophic plants growing in forests - mistletoes and partial mycoheterotrophs. It is remarkable that despite apparent evolutionary pressure towards improved carbon acquisition from the host, M. pratense retains efficient photosynthesis and high transpiration rate, the ecophysiological traits typical of related root hemiparasites in the Orobanchaceae.

Ultrastructure of hydathode trichomes of hemiparasitic Rhinanthus alectorolophus and Odontites vernus: how important is their role in physiology and evolution of parasitism in Orobanchaceae?

The Rhinanthoid clade of the family Orobanchaceae comprises plants displaying a hemiparasitic or holoparasitic strategy of resource acquisition. Some of its species (mainly Rhinanthus spp.) are often used as models for studies of hemiparasite physiology. Although there is a well-developed concept covering their physiological processes, most recent studies have neglected the existence of hydathode trichomes present on leaves of these hemiparasitic plants. As a first step for the proposed integration of these structures in the theory of physiological processes of the hemiparasites, we described the outer micromorphology and ultrastructure of the hydathode trichomes on leaves of hemiparasitic Rhinanthus alectorolophus and Odontites vernus with scanning and transmission electron microscopy (SEM and TEM, respectively). The TEM inspections of both types of trichome revealed typical ultrastructural features: labyrinthine cell wall, high content of cytoplasm in cells with numerous mitochondria and presence of plasmodesmata. All these features indicate high metabolic activity complying with their function as glandular trichomes actively secreting water. The active secretion of water by the hydathode trichomes (evidence for which is summarised here) also presents a possible mechanism explaining results of previous gas exchange measurements detecting high dark respiration and transpiration rates and a tight inter-correlation between them in hemiparasitic Orobanchaceae. In addition, this process is hypothesised to have allowed multiple evolutionary transitions from facultative to obligate hemiparasitism and unique xylem-feeding holoparasitism of Lathraea with a long-lived underground stage featuring a rhizome covered by scales of leaf origin.