Herbs are not just small plants: What biomass allocation to rhizomes tells us about differences between trees and herbs.

PREMISE: Biomass accumulation over years in vertical stems of trees leads to hypoallometric scaling between stem and leaf biomass within this growth form, while for herbaceous species, biomass allocation between these organ types typically exhibits isometry. However, biomass accumulation in herbs can occur in belowground perennating organs (e.g., rhizomes) that are, contrary to aboveground parts of herbs, long-lived. Although ecologically important, biomass allocation and accumulation in rhizomes (and similar organs) have mostly not been studied. METHODS: We assembled data on biomass investments into plant organs for 111 rhizomatous herbs based on a literature survey and greenhouse experiment. We estimated the proportion of whole-plant biomass invested into rhizomes and, using allometric relationships, analyzed scaling between rhizome and leaf biomass and whether it is more variable than for other organs. RESULTS: On average, rhizomes comprise 30.2% of the total plant biomass. The proportion allocated to rhizomes does not change with plant size. Scaling between rhizome and leaf biomass is isometric, and allocation to rhizomes is not more variable than allocation to other organs. CONCLUSIONS: Rhizomatous herbs accumulate substantial biomass in rhizomes, and rhizome biomass scales isometrically with leaves, contrary to the hypoallometric relationship between stem and leaves in trees. This difference suggests that the rhizome biomass is in balance with aboveground biomass-a resource of carbon for rhizome formation that, at the same time, is dependent on carbon stored in rhizomes for its seasonal regrowth.

Genome size of alpine plants does not predict temperature resistance.

MAIN CONCLUSION: Genome size of alpine plants is not related to their resistance against frost and heat. Genome size is a variable trait in angiosperms, and it was suggested that large genome size represents a constraint in stressful environments. We measured genome size and resistance to frost and heat in 89 species of plants from tropical and temperate alpine habitats. Genome size of the species, ranging from 0.49 pg to 25.8 pg across the entire dataset, was not related to either frost or heat resistance in either group of plants. Genome size does not predict resistance to extreme temperatures in alpine plants and is thus not likely to predict plant responses to climate changes.

Comparative anatomy of leaf petioles in temperate trees and shrubs: the role of plant size, environment and phylogeny.

BACKGROUND AND AIMS: Petioles are important plant organs connecting stems with leaf blades and affecting light-harvesting ability of the leaf as well as transport of water, nutrients and biochemical signals. Despite the high diversity in petiole size, shape and anatomy, little information is available regarding their structural adaptations across evolutionary lineages and environmental conditions. To fill this knowledge gap, we investigated the variation of petiole morphology and anatomy of mainly European woody species to better understand the drivers of internal and external constraints in an evolutionary context. METHODS: We studied how petiole anatomical features differed according to whole-plant size, leaf traits, thermal and hydrological conditions, and taxonomic origin in 95 shrubs and trees using phylogenetic distance-based generalized least squares models. KEY RESULTS: Two major axes of variation were related to leaf area and plant size. Larger and softer leaves are found in taller trees of more productive habitats. Their petioles are longer, with a circular outline and are anatomically characterized by the predominance of sclerenchyma, larger vessels, interfascicular areas with fibres and indistinct phloem rays. In contrast, smaller and tougher leaves are found in shorter trees and shrubs of colder or drier habitats. Their petioles have a terete outline, phloem composed of small cells and radially arranged vessels, fibreless xylem and lamellar collenchyma. Individual anatomical traits were linked to different internal and external drivers. Petiole length and vessel diameter increase with increasing leaf blade area. Collenchyma becomes absent with increasing temperature, and petiole outline becomes polygonal with increasing precipitation. CONCLUSIONS: We conclude that species' temperature and precipitation optima, plant height, and leaf area and thickness exerted a significant control on petiole anatomical and morphological structures not confounded by phylogenetic inertia. Species with different evolutionary histories but similar thermal and hydrological requirements have converged to similar petiole anatomical structures.

Carbohydrate storage in herbs: the forgotten functional dimension of the plant economic spectrum.

BACKGROUND AND AIMS: Although the plant economic spectrum seeks to explain resource allocation strategies, carbohydrate storage is often omitted. Belowground storage organs are the centre of herb perennation, yet little is known about the role of their turnover, anatomy and carbohydrate storage in relation to the aboveground economic spectrum. METHODS: We collected aboveground traits associated with the economic spectrum, storage organ turnover traits, storage organ inner structure traits and storage carbohydrate concentrations for ~80 temperate meadow species. KEY RESULTS: The suites of belowground traits were largely independent of one another, but there was significant correlation of the aboveground traits with both inner structure and storage carbohydrates. Anatomical traits diverged according to leaf nitrogen concentration on the one hand and vessel area and dry matter content on the other; carbohydrates separated along gradients of leaf nitrogen concentration and plant height. CONCLUSIONS: Contrary to our expectations, aboveground traits and not storage organ turnover were correlated with anatomy and storage carbohydrates. Belowground traits associated with the aboveground economic spectrum also did not fall clearly within the fast-slow economic continuum, thus indicating the presence of a more complicated economic space. Our study implies that the generally overlooked role of storage within the plant economic spectrum represents an important dimension of plant strategy.

Young clonal and non-clonal herbs differ in growth strategy but not in aboveground biomass compensation after disturbance.

Clonal plants have more traits enabling individual persistence (larger belowground storage of buds and assimilates), whereas non-clonal plants have more traits enabling population persistence (a higher reliance on regeneration from seeds). This difference presumably makes those groups respond differently to disturbance. We asked whether this difference is already expressed in the first year of the plant's life. In a pot experiment with 17 congeneric pairs of clonal and non-clonal herbs, we investigated response to a disturbance at the individual level. We were interested whether the leaf C/N ratio (a proxy reflecting active growth and photosynthetic efficiency), the R/S ratio (a proxy for belowground storage) and the amount of compensated biomass differ between clonal and non-clonal herbs. Moreover, we asked whether compensation for the loss of aboveground biomass after disturbance can be predicted by the R/S ratio or explained by the leaf C/N ratio. We found that clonal herbs have higher leaf C/N and R/S ratios than non-clonal herbs. Under disturbance, the leaf C/N and R/S ratios decreased in the clonal herbs and increased in the non-clonal herbs. However, the clonal and non-clonal plants did not differ in biomass compensation ability. Neither the R/S ratio nor the leaf C/N ratio explained the compensation abilities of the herbs. These results show that even though the growth strategies of clonal and non-clonal plants and their reactions to disturbance are different, the groups are similarly capable of compensating for the loss of aboveground biomass. Clonal plants do not have an advantage over non-clonal plants under disturbance during their first year of life.

The Effect of Rhizoboxes on Plant Growth and Root: Shoot Biomass Partitioning.

Various types of flat rhizoboxes aid in root visualization and tracking in experiments where the focus is upon root system growth and development. While size of the pot is known to affect experiments, nothing is known about the impact of rhizoboxes-not only their volume, but also their shape might affect root and shoot growth. Therefore, we investigated how rhizoboxes change plant biomass and root:shoot biomass partitioning. We compared biomass and root:shoot ratio of plants growing in the pots with different geometry-usual three-dimensional, cuboid plant pots and flat two-dimensional rhizoboxes about the same volume. We used two different nutritional treatments (deionized water and additional nutrients) for investigating whether the nutrient availability in the substrate changed the impact of rhizoboxes on plant growth. We used 15 species for the generalizability of our results across the phylogenetic tree. Proportional investment of plants into roots was similar in usual pots and in rhizoboxes. This pattern was stable across nutrition treatments and across species. Further, we found no differences in total biomass of plants between pot type within nutrient treatments. With added nutrients, the plants had a higher biomass and lower root:shoot ratio compared to treatments without nutrient addition. Thus, species can be safely compared when grown in the rhizoboxes; rhizoboxes did not affect root system growth comparisons among species and nutrient levels. Also, they did not affect plant growth in terms of total biomass.