Introduction

Julie L. Whitbeck and Zoe G. Cardon

Below the soil surface, the rhizosphere is the crossroads of the soil habitat,
a hub of biological, chemical, and physical activity surrounding the living
infrastructure of plant roots. Complex fine-scale gradients of substrate availability, water potential, and redox state distinguish this habitat from bulk soil
and constrain the distribution and the activity of the tremendously diverse
rhizosphere biota. Populations of archea, bacteria, protists, fungi, and animals
live here along with plant roots, the activities of each influencing those of
the others across spatial and temporal scales spanning orders of magnitude.
The nature of the exchange and transformation of universal biological currencies - resources such as organic carbon, mineral nutrients and water - by
these biota determines paths of energy flow and shapes community structure
and ecosystem properties. Information is also exchanged among rhizosphere
inhabitants, via mechanisms including quorum sensing and the production
of phytohormone mimics. The influence of rhizosphere activity extends far
beyond the rhizosphere itself, manifest across the landscape and through time
in patterns of community structure and ecosystem processes, and in patterns
of soil development.
Although understanding of soil biological, physical, and chemical function
has lagged behind comprehension of aboveground processes, insight into
belowground function is essential for grappling with current environmental
challenges in natural and managed terrestrial ecosystems worldwide. Interest
in rhizosphere ecology has a long history with roots in agronomy, mycology,
plant physiology, and microbiology. Contemporary study of the rhizosphere
is, by necessity, interdisciplinary, depending on understanding soil physical
and chemical properties, plant biology, and the activity and organization of
microbes and soil fauna. Syntheses focusing on the rhizosphere (e.g. Fitter
1985; Box Jr. and Hammond 1990; Lynch 1990) reflect this interdisciplinary
nature and also note the applied value of rhizosphere research for natural and
agricultural ecosystem management. Several more recent offerings have delved
into specialized areas of rhizosphere biology, often employing reductionist
approaches to examine the nature and function of specific kinds of interactions
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Introduction

(e.g. rhizosphere biochemistry in Pinto etal. 2001, solute transport in Tinker

and Nye 2000, biogeochemistry of trace elements in Gobran etal 2001, and
Huang and Gobran 2005), or addressing rhizosphere management in the
context of particular goals (e.g. Wright and Zobel 2005).
Within the field of ecology, attention to the rhizosphere has grown extensively and rapidly since the mid-1980s, spanning the full breadth of biological
and biogeochemical inquiry from ephemeral shifts in bacterial enzyme production in microliter volumes of soil to landscape scale dynamics of soil genesis
over millenia. This book models its cross-scale and interdisciplinary approach
after Fitter's 1985 edited volume Ecological Interactions in Soil, updating our
ecological frame of reference for the rhizosphere. Our goal is to invigorate
interaction among scientists working on diverse aspects of rhizosphere ecology and to pique the interest of a broad audience interested generally in
belowground ecological function in terrestrial ecosystems. Contributions from
a range of scientists focus on rhizosphere ecology and the emergent consequences of rhizosphere activity, including chapters addressing soil biota (plant
roots, microbes, soil fauna), interactions among organisms and soils, and the
implications of those interactions for rhizosphere trophic organization, productivity, nutrient cycling, soil genesis, and ecosystem management. Instead
of writing reviews, authors present perspectives on the rhizosphere, including key developments and the interdisciplinary or cross-scale connections
linking their focal research area into the network of ecological rhizosphere
research. Complementary views of the rhizosphere from very different spatial
and temporal perspectives, and at varied levels of abstraction, overlap in at
least three major areas detailed below: rhizosphere soil biogeochemistry and
physical structure, taxonomic and functional diversity of rhizosphere biota,
and integration and coordination of rhizosphere interactions.
First, a better understanding of the biogeochemical and physical nature of
the rhizosphere environment and habitat and a corresponding insight into the
influence of rhizosphere ecology on the trajectory of soil genesis emerge from
several chapters. Richter et al. (Chapter 8) highlight the central role played by
roots and associated rhizospheres in chemical and physical weathering, positing that over pedogenic timescales, rhizospheres are fundamental drivers of
dramatic soil biogeochemical transformation and overall soil development, so
fundamental that almost all soil might be viewed as rhizosphere soil of varying
age. Hawkes etal. (Chapter 1) begin their chapter with the same notion, then
delve into the small-scale yet dramatic redox (Richter etal.), resource (Cheng
and Gershenson, Chapter 2), water, and other gradients surrounding single
plant roots that directly affect microbial community functions such as nitrogen
cycling. Such small-scale gradients around roots are illustrated, for example,
by rhizosphere-induced soil mottling (Richter etal.), and their presence has
tremendous implications for ecosystem responses to global change (Cheng

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XVU

and Gershenson, Chapter 2; Pregitzer etal. Chapter 7) when increased [COj]

and N-deposition induce shifts in fine root turnover, extent of rooting, root
respiration, and associated rhizosphere microbial community composition and
activity.
Soil physical structure is also influenced strongly by rhizosphere biota and
their activities, again with implications from microbial to ecosystem scales. Soil
aggregates essential for maintaining soil porosity and conductivity, housing
microsites for denitrification, and protecting soil carbon from decomposition
are bound and stabilized by fungal hyphae and bacterial exopolysaccharides
(Drinkwater and Snapp, Chapter 6; Johnson and Gehring, Chapter 4). Plant
roots can lift and mix surface soil layers over generations; great increases in
rhizosphere bulk density (and associated decreases in conductivity and pore
space) can be driven by single roots as they expand in diameter in deeper soil
horizons (Richter et al.. Chapter 8). Even the slow, physical breakdown of rock
is facilitated by generations of roots as they penetrate channels and generate
fractures (Richter etal). The physical structure and organization of rhizospheres themselves, and their persistent imprints on soil physical structure,
are emerging as potential key controllers of biogeochemistry and biological
interaction across spatial and temporal scales (e.g. Crawford etal. 2005).
A second theme is the consideration of patterns and consequences of taxonomic and functional diversity in the rhizosphere. For example, even at
global scales, functional biodiversity aligned with evolutionary lineage among
mycorrhizae has great implications for large, biogeographic patterns in belowground function (Johnson and Gehring, Chapter 4); saprophytic capabilities
are minimal among arbuscular mycorrhizae and maximal among cricoid mycorrhizae, suggesting a biogeographic gradient from grasslands to tundra in
the reliance of the mycorrhizal symbiotic partners on free-living saprotrophs
for mineralization of nutrients from organic matter. Drinkwater and Snapp
(Chapter 6) emphasize the importance of re-establishing biodiversity within
the rhizosphere in order to redevelop self-sustaining agricultural systems that
require reduced fertilizer inputs. They underscore the idea that, prior to the
heavy-input, mechanized agriculture prevalent today, plants and associated
rhizosphere biota evolved together within the functioning rhizosphere system,
only to be separated conceptually and actually when tillage, fertilizer, and
pest control inputs were implemented. These large spatial and long temporal
views of patterns in rhizosphere communities are complemented by Hawkes
etal and Griffiths etal (Chapters 1 and 3) who focus explicitly on specific
community membership and signaling. Communities of rhizosphere microbes
and soil fauna shift not only with soil management techniques but also as a
function of plant species and soil type (Garbeva etal 2004; Griffiths etal,
Chapter 3; Hawkes etal. Chapter 1), yet the implications of diversity for

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resilience of the soil community and for maintenance of ecosystem functions

remain unknown.
The third theme is the quest to understand controls over integrative balance and versatile coordination in the rhizosphere across scales of biological
organization. For example, Moore etal. (Chapter 5) suggest that the fundamental structure of rhizosphere food webs, with multiple (bacterial, fungal,
and root) channels through which energy can flow to higher trophic levels,
supports web stability even in the face of shifting food-web membership on
ecological, and evolutionary, timescales. It is general community structure,
not the specific community members, that is most important in capturing the
essence of rhizosphere community energetic function and stability. Hawkes
et al. (Chapter 1) and Griffiths et a\. (Chapter 3) suggest that, beyond such generalized trophic relationships, specific chemical communication among diverse
rhizosphere community members is a key determinant of ecological function.
Notable mechanisms include a plethora of signaling molecules that enable
communication and activity coordination among microbes themselves (e.g.
quorum sensing signals) and among microbes and plant roots (e.g. molecules
similar to plant hormones produced by microbes or by microfauna). Again,
in Chapter 6, Drinkwater and Snapp address implications of this flexibility in
community composition for agroecosystem management, while in Chapter 7,
Pregitzer etal. query how resilient rhizosphere community structure will be
to global scale shifts in carbon and nitrogen availability.
These contrasting and complementary views suggest that the most powerful
insights into the essence of rhizosphere ecology will grow from the synergy
of reductionist and integrative ecosystems approaches. Since the organismal
diversity in the rhizosphere is enormous, and the suite of potential interactions and mechanistic controllers is too large for all to be examined in detail,
broader scale properties, biogeochemical or biogeographical setting, or historical background can be used to help guide the focus and interpretation
of mechanistic investigations. The potential implications of newly discovered
fine-scale rhizosphere patterns and mechanisms can then be more readily
considered in work addressing larger-scale or higher-order ecological system function. Focusing on just one of many promising research directions,
recent advances illuminating the nature of the rhizosphere habitat at quite
fine-scale resolution, along with a growing appreciation of the relevance of
rhizosphere ecological activity for long-term soil development and in the service of human needs, provide the basis for designing studies to investigate the
kinds, complexity, and extent of the feedbacks between soil physical structure
and ecological processes in the rhizosphere. For example, research examining
rhizosphere bacterial physiological responses to variation in soil aggregation
can be strengthened by understanding the temporal and spatial scales of variation in soil properties and the ecosystem processes to which these organisms

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XIX

contribute. Likewise, ecosystem scale investigations of soil carbon sequestration can draw upon understanding of the key physiological and community
level properties that control carbon and energy flow in the rhizosphere, in
order to link changes in soil structure with changes in carbon content over
time and/or across landscapes.
From our perspectives as rhizosphere ecologists, we hope the contributions
to this book inspire research that draws upon cross-scale and interdisciplinary
understanding to develop new insights into rhizosphere ecology and management, as well as into ecology as a whole.

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