Internal transport of gases is crucial for vascular plants inhabiting aquatic, wetland or flood-prone environments. Diffusivity of gases in water is approximately 10 000 times slower than in air; thus direct exchange of gases between submerged tissues and the environment is strongly impeded. Aerenchyma provides a low-resistance internal pathway for gas transport between shoot and root extremities. By this pathway, O2 is supplied to the roots and rhizosphere, while CO2, ethylene, and methane move from the soil to the shoots and atmosphere. Diffusion is the mechanism by which gases move within roots of all plant species, but significant pressurized through-flow occurs in stems and rhizomes of several emergent and floating-leaved wetland plants. Through-flows can raise O2 concentrations in the rhizomes close to ambient levels. In general, rates of flow are determined by plant characteristics such as capacity to generate positive pressures in shoot tissues, and resistance to flow in the aerenchyma, as well as environmental conditions affecting leaf-to-air gradients in humidity and temperature. O2 diffusion in roots is influenced by anatomical, morphological and physiological characteristics, and environmental conditions. Roots of many (but not all) wetland species contain large volumes of aerenchyma (e.g. root porosity can reach 55%), while a barrier impermeable to radial O2 loss (ROL) often occurs in basal zones. These traits act synergistically to enhance the amount of O2 diffusing to the root apex and enable the development of an aerobic rhizosphere around the root tip, which enhances root penetration into anaerobic substrates. The barrier to ROL in roots of some species is induced by growth in stagnant conditions, whereas it is constitutive in others. An inducible change in the resistance to O2 across the hypodermis/exodermis is hypothesized to be of adaptive significance to plants inhabiting transiently waterlogged soils. Knowledge on the anatomical basis of the barrier to ROL in various species is scant. Nevertheless, it has been suggested that the barrier may also impede influx of: (i) soil-derived gases, such as CO2, methane, and ethylene; (ii) potentially toxic substances (e.g. reduced metal ions) often present in waterlogged soils; and (iii) nutrients and water. Lateral roots, that remain permeable to O2, may be the main surface for exchange of substances between the roots and rhizosphere in wetland species. Further work is required to determine whether diversity in structure and function in roots of wetland species can be related to various niche habitats.

https://onlinelibrary.wiley.com/doi/pdf/10.1046/j.1365-3040.2003.00846.x

Long-distance transport of gases in plants: a perspective on internal aeration and radial oxygen loss from roots

Life on Earth is sustained by a small volume of soil surrounding roots, called the rhizosphere. The soil is where most of the biodiversity on Earth exists, and the rhizosphere probably represents the most dynamic habitat on Earth; and certainly is the most important zone in terms of defining the quality and quantity of the Human terrestrial food resource. Despite its central importance to all life, we know very little about rhizosphere functioning, and have an extraordinary ignorance about how best we can manipulate it to our advantage. A major issue in research on rhizosphere processes is the intimate connection between the biology, physics and chemistry of the system which exhibits astonishing spatial and temporal heterogeneities. This review considers the unique biophysical and biogeochemical properties of the rhizosphere and draws some connections between them. Particular emphasis is put on how underlying processes affect rhizosphere ecology, to generate highly heterogeneous microenvironments. Rhizosphere ecology is driven by a combination of the physical architecture of the soil matrix, coupled with the spatial and temporal distribution of rhizodeposits, protons, gases, and the role of roots as sinks for water and nutrients. Consequences for plant growth and whole-system ecology are considered. The first sections address the physical architecture and soil strength of the rhizosphere, drawing their relationship with key functions such as the movement and storage of elements and water as well as the ability of roots to explore the soil and the definition of diverse habitats for soil microorganisms. The distribution of water and its accessibility in the rhizosphere is considered in detail, with a special emphasis on spatial and temporal dynamics and heterogeneities. The physical architecture and water content play a key role in determining the biogeochemical ambience of the rhizosphere, via their effect on partial pressures of O2 and CO2, and thereby on redox potential and pH of the rhizosphere, respectively. We address the various mechanisms by which roots and associated microorganisms alter these major drivers of soil biogeochemistry. Finally, we consider the distribution of nutrients, their accessibility in the rhizosphere, and their functional relevance for plant and microbial ecology. Gradients of nutrients in the rhizosphere, and their spatial patterns or temporal dynamics are discussed in the light of current knowledge of rhizosphere biophysics and biogeochemistry. Priorities for future research are identified as well as new methodological developments which might help to advance a comprehensive understanding of the co-occurring processes in the rhizosphere.

Rhizosphere: biophysics, biogeochemistry and ecological relevance

Roots of virtually all vascular plants have an endodermis with a Casparian band, and the majority of angiosperm roots tested also have an exodermis with a Casparian band. Both the endodermis and exodermis may develop suberin lamellae and thick, tertiary walls. Each of these wall modifications has its own function(s). The endodermal Casparian band prevents the unimpeded movement of apoplastic substances into the stele and also prevents the backflow of ions that have moved into the stele symplastically and then were released into its apoplast. In roots with a mature exodermis, the barrier to apoplastic inflow of ions occurs near the root surface, but prevention of backflow of ions from the stele remains a function of the endodermis. The suberin lamellae protect against pathogen invasion and possibly root drying during times of stress. Tertiary walls of the endodermis and exodermis are believed to function in mechanical support of the root, but this idea remains to be tested. During stress, root growth rates decline, and the endodermis and exodermis develop closer to the root tip. In two cases, stress is known to induce the formation of an exodermis, and in several other cases to accelerate the development of both the exodermis and endodermis. The responses of the endodermis and exodermis to drought, exposure to moist air, flooding, salinity, ion deficiency, acidity, and mechanical impedance are discussed.

Root Endodermis and Exodermis: Structure, Function, and Responses to the Environment

Anoxia can be one consequence of waterlogging and submergence of plants. Anoxia in plant tissues reduces the rate of energy production by 65-97% compared with the rate in air. Thus, adaptation to anoxia always includes coping with an energy crisis. Tolerance to anoxia is relevant to wetland species, rice cultivation and transient waterlogging of other agricultural and horticultural crops. This perspective, in two parts, examines mechanisms of anoxia tolerance in plants. Part 1 covers anoxia tolerance in terms of growth and survival, the interaction of anoxia tolerance with other environmental factors, and the development of anoxic cores within plant tissues. Equally importantly, Part 1 also examines anaerobic carbohydrate catabolism (principally ethanolic fermentation in plants) and its regulation. We put forward two modes of anoxia tolerance, one based on reduced rates of anaerobic carbohydrate catabolism and the other on accelerated rates (Pasteur effect). Further, Part 1 examines mechanisms of post-anoxic injury. In Part 2 (Greenway and Gibbs, manuscript in preparation) we consider flow of the limited amount of energy produced under anoxia to processes essential for cell survival. We show that acclimation to anoxia in plants involves integration of a set of sophisticated characteristics, as a consequence of which the habitat within the anoxic cell is a very different world to that of the aerobic cell.

Review: Mechanisms of anoxia tolerance in plants. I. Growth, survival and anaerobic catabolism

Biodiversity is a challenging field of research. Approaches are manifold and mostly cover few aspects of the total wealth of phenomena only. The mapping of the vascular plants and the inventory of Basidiomycota in Bavaria are projects to be mentioned in this context as one part of the commitment of the author. In the following article the author describes further activities during his lifework in regard A. Bresinsky (*) Institute of Botany of University of Regensburg, D-93040, Regensburg, Germany e-mail: abresinsky@t-online.de U. Lüttge et al. (eds.), Progress in Botany, Progress in Botany 75, DOI 10.1007/978-3-642-38797-5_1, © Springer-Verlag Berlin Heidelberg 2014 3 to ant–plant interactions resulting in dispersal and distribution of plants, and moreover, in respect to speciation and to evolution of high ranked taxa within the fungi, discussed mainly in relation to the genus Pleurotus and the order Boletales. In fungi the investigations include breeding systems, isolation barriers, polyploidy, pigment patterns and DNA phylogeny. The pigment patterning in Boletales correlates well with the phylogeny as revealed by DNA analysis of selected gene sections.

Progress in Botany

Oxygen Distribution in Wetland Plant Roots and Permeability Barriers to Gas-exchange with the Rhizosphere: a Microelectrode and Modelling Study with Phragmites australis

P. M. Beckett

Papers

Oxygen Distribution in Wetland Plant Roots and Permeability Barriers to Gas-exchange with the Rhizosphere: a Microelectrode and Modelling Study with Phragmites australis