ARTICLE IN PRESS

Soil Biology & Biochemistry 38 (2006) 31813183

[Link]/locate/soilbio

Editorial

Ecosystems in ux: Molecular and stable isotope assessments of soil

organic matter storage and dynamics
Soil organic carbon is the largest pool (1500 Pg C
(1 Pg 1015 g)) in the terrestrial carbon cycle, and contains
twice as much C as the atmosphere (780 Pg C) and nearly
triple the amount of C in terrestrial biomass (550 Pg C)
(Schlesinger, 1997; Amundson, 2001; Houghton, 2005).
The soil organic carbon (SOC) pool is also a dynamic
component of the C cycle that is closely linked to the
atmospheric CO2 pool via inputs from dead organic matter
production (E60 Pg C yr
1) and losses from decomposition
(E60 Pg C yr
1) (Schlesinger, 1997; Houghton, 2005).
These uxes are 10-fold greater than fossil fuel combustion
(E6 Pg C yr
1), so that even small changes in the magnitude of the SOC pool or the input/output rates associated
with it could have profound consequences for the global C
cycle, the concentration of CO2 in the atmosphere, and the
climate system (Lal et al., 1995).
Despite the obvious signicance of SOC, our present
concept of the global C cycle remains limited by
uncertainties in the quantitative aspects of SOC storage
and dynamics. On the one hand, historical reconstructions
of land-use activities (agriculture and deforestation in
particular) reveal that the global SOC pool has decreased
by 5578 Pg C since AD 1850, implying that soils have been
a signicant source of atmospheric CO2 in recent history
(Lal et al., 1995; Lal, 2004). On the other hand, indirect
estimates based on geophysical constraints and inverse
modeling of the atmosphere and oceans indicate that
terrestrial ecosystems and soils must be presently accumulating C (Fan et al., 1998; Houghton, 2005). To reconcile
these differences, we must enhance our understanding of
the quantitative details and mechanisms of soil organic
matter (SOM) storage and dynamics in relation to changes
in land cover/land use, climate, and the gas composition of
the atmosphere (Bellamy et al., 2005; Fung et al., 2005;
Heath et al., 2005; Jastrow et al., 2005; Potter et al., 2006).
This goal permeates nationally and globally identied
scientic needs and is essential to be able to consider
sustainable management of the Earth system (NRC,
2001b).
In addition to its role in the function of the Earth system
at the global scale, SOM also plays a pivotal role in the
provision of ecosystem services that are essential to the
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well-being of the human population (Daily et al., 1997;

Millenium Ecosystem Assessment, 2005). These key
ecosystem services include the maintenance of soil fertility,
nutrient cycling, production of food and ber, inltration
and storage of water, erosion and ood control, and the
maintenance of biodiversity. August scientic bodies such
as the National Academy of Sciences of the USA,
International Union of Soil Sciences, US Carbon Cycle
Program, the United Nations Millenium Ecosystem
Assessment, and the Intergovernmental Panel on Climate
Change all rate the mechanisms and processes that control
SOM storage and dynamics among the most fundamental
areas of science essential for an understanding of the Earth
system. For example, of the eight Grand Challenges in
environmental sciences identied by the US National
Research Council that require enhanced scientic understanding and federal research funding, four of them (i.e.
biogeochemical cycles, biological diversity and ecosystem
functioning, climate variability, and land-use dynamics)
have SOM dynamics as either an explicit or implicit central
issue (NRC, 2001a). Additionally, the US NRC in their
publication on basic research opportunities in the Earth
science (NRC, 2001b) also places a particularly important
emphasis on the need for greater understanding of the
processes determining how soils control and respond to
changes in the Earth system.
Over the last decade, a multidisciplinary collection of
scientists have converged on this important issue, bringing
fundamental knowledge in pedology, mineralogy, biogeochemistry, the biology of plants and microbes, and isotope
chemistry to bear on this problem. To highlight recent
advances resulting from these collaborations, we convened
a symposium entitled Ecosystems in Flux: Molecular and
Stable Isotope Assessments of Soil Organic Matter Storage
and Dynamics held at the Fall 2004 American Geophysical Union (Biogeosciences Division) in San Francisco,
California, USA. This session brought together scientists
from many disciplines to discuss the interactions between
soil structure, mineralogy, organic chemistry, and microbial community structure and activity, and how these
factors determine the storage and turnover of organic C
and nitrogen (N) in managed and natural ecosystems. In

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Editorial / Soil Biology & Biochemistry 38 (2006) 31813183

total, 42 presentations (28 posters and 14 talks) were given

by scientists from Europe and the United States on topics
including the chemical, physical and structural mechanisms
of SOM stabilization among various ecosystems, the
accessibility of C pools to soil microbes, the specic role
of microbial communities and their enzyme systems in
SOM dynamics, and the response of SOM pools to
perturbations such as land uses, land cover changes, and
rising atmospheric CO2 concentration. This special issue of
Soil Biology and Biochemistry represents some of the key
contributions to that session.
This volume contains four contributions with the goal of
directly or indirectly tracking the role of microbial activity
on SOM dynamics through studies of enzyme activity, the
isotopic composition of respired CO2, and the isotopic
composition of phospholipids and extractable microbial
biomass. The paper by Kramer and Gleixner investigated
the relative utilization of fresh particulate vs. old SOM by
Gram-negative and Gram-positive bacteria in agricultural
soils through combined 14C and 13C compound specic
isotope analysis of phospholipids. Allison and Jastrow used
physical fractionation procedures to investigate whether
soil C is spatially isolated from degradative enzymes across
a prairie restoration chronosequence, and found that
enzyme production and C turnover occurred rapidly in
particulate organic matter (POM) fractions, but slowly in
mineral-dominated fractions where enzymes and their C
substrates were immobilized on mineral surfaces. Dijkstra
et al. presented a new technique for the isolation of
microbial biomass for isotope analysis, and documented an
enrichment in the natural abundances of 13C and 15N in
microbial biomass relative to SOM across a broad range of
soil types and vegetation. These results reected the extent
to which microbial products contributed to stabilized pools
of SOM, and helped explain the 13C and 15N enrichment
that is commonly observed along the SOM decay
continuum. Crow et al. used density fractionation to
separate light POM fractions from heavy, mineral-associated SOM fractions, and examined the accessibility of
these pools to microbial inoculants by quantifying the
amount and d13C value of CO2 respired from those SOM
density fractions.
Four additional papers illustrated the importance of
land use and land cover in controlling SOM dynamics. In
two of these manuscripts, Liao et al. elucidated the
mechanisms responsible for changes in SOC and soil total
N apportionment among aggregated and non-aggregated
mineral-associated and POM following woody plant
encroachment into grassland. To accomplish this, they
analyzed the elemental and stable isotope (d13C and d15N)
abundances of specic size/density fractions along a
chronosequence of C3 woody plant encroachment into
C4 grassland. The paper by Krull et al. employed
compound-specic d2H and d13C analyses of n-alkanes
from SOM in a grasslandwoodland system to examine
interactions between carbon and water cycles, and to
evaluate the two-layer soil water model in a semi-arid

savanna system. Yamishita et al. analyzed the effect of land

cover/land use (spruce forest, grassland, wheat and maize)
on the distribution and turnover of SOM in size/density
fractions. The mass and C content of both the free and
occluded POM fractions were greater in the forest soil than
in the grassland and arable soils, and 13C kinetics indicated
that the free POM fractions had the most rapid rates of
turnover.
The effects of rising CO2 concentrations on SOC
allocation and subsequent SOM dynamics has garnered
signicant attention recently, and is the focus of the paper
by Del Galdo et al. They quantied SOC stores and
dynamics in relationship to soil aggregation and pool
composition in a Californian chaparral ecosystem exposed
for 6 years to a gradient of atmospheric CO2. They found
that increasing atmospheric CO2 concentrations decreased
soil C in chaparral, and that the microaggregate fraction
was the most responsive to elevated concentrations of
atmospheric CO2.
Soil organic N dynamics are extremely important for
forest, grassland, and agroecosystem productivity studies,
but the state of knowledge of the mechanistic controls on N
pools and the structural composition of stabilized pools is
insufcient. Three manuscripts in this volume address these
issues. Sollins et al. explored the common observation that
C concentrations and C/N generally decrease with particle
density. They applied detailed chemical, isotopic, molecular, and mineralogical analyses to a density-fractionated
andic soil. Results were consistent with a general pattern of
an increase in the extent of microbial processing with
increasing organo-mineral particle density, and also with
an onion layering model of SOM accumulation on
mineral surfaces in which polar carboxyl and nitrogenous
compounds sorb by ligand and cation exchange. The
manuscript by Olk et al. examined yield decreases in several
agricultural systems associated with repeated cropping
under wet or submerged soil conditions, and attributed this
decline in yield to the accumulation of phenolic lignin
compounds that covalently bind nitrogenous compounds
into recalcitrant forms. Recent advances in 15N-NMR
allowed them to identify agronomically signicant quantities of lignin-bound N in rice cropping systems. Kolbl et
al. applied 15N-labelled mustard litter to an agricultural
system to examine N-cycling processes in low yield vs. high
yield portions of the landscape. The distribution of 15N was
followed in POM fractions and in ne mineral fractions.
They found that the higher silt and clay contents of low
yield areas promoted N stabilization in ne mineral
fractions.
In total, the scientic contributions of these papers were
many and varied, spanning a broad range of important
SOM-related issues within climate change, land use,
microbiology, soil chemistry, and soil structure. Furthermore, several of these papers present new and innovative
technical advances in the use of isotopic and molecular
techniques in soil science. These contributions will better
enable the scientic community to understand and predict

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Editorial / Soil Biology & Biochemistry 38 (2006) 31813183

patterns and processes in the soil environment at scales

ranging from the molecular to the global level. We hope
these contributions will stimulate and inuence further
research into this fundamentally important area of Earth
system science.
Acknowledgements
As guest editors, we thank the authors who contributed to
this special issue as well as to all of those who presented talks
and posters at the AGU meeting. We are especially grateful
to the many colleagues who contributed signicant time and
effort to the review process: Wulf Amelung (University of
Bayreuth), Ron Amundson (University of California, Berkeley), Jeff Baldock (CSIRO Land and Water), Bruce Caldwell
(Oregon State University), Susan Crow (Oregon State
University), Xiaoyan Dai (Texas A&M University), MarieFrance Dignac (University of Paris), Karl Dria (Purdue
University); Ryan Fimmen (The Ohio State University), Alan
Franzluebbers (United States Department of AgricultureARS), Rick Gill (Washington State University), Miquel
Gonzalez-Meler (University of Illinois, Chicago), Roland Bol
(Institute of Grassland and Environmental Research), Yongsong Huang (Brown University), Erik Hobbie (University of
New Hampshire), Rob Jackson (Duke University), Julia Liao
(Rice University), Hans-Dietrich Luedemann (University of
Regensburg), Carrie Masiello (Rice University), David
Myrold (Oregon State University), Mark Pagani (Yale
University), Elise Pendall (University of Wyoming), Martin
Potthoff (Georg August University of Gottingen), Peter
Sauer (Indiana University), Alex Sessions (California Institute of Technology), Josh Schimel (University of California, Santa Barbara), Kate Scow (University of California,
Davis), Ron Smernik (University of Adelaide), Phil Sollins
(Oregon State University), Diane Stott (United States
Department of Agriculture-ARS), and Susan Trumbore
(University of California, Irvine). Additionally, we thank
John Waid, Chief Editor, Soil Biology & Biochemistry as well
as the editorial staff at Elsevier.
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T.R. Filley
Department of Earth and Atmospheric Sciences, Purdue
University, West Lafayette, IN 47907-1397, USA
E-mail address: lley@[Link]
T.W. Boutton
Department of Rangeland Ecology and Management, Texas
A&M University, College Station, TX 77843-2126, USA
E-mail address: boutton@[Link]