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Design of reserve networks and the

persistence of biodiversity
Mar Cabeza and Atte Moilanen

Sophisticated computational methods have been developed to help us to problem: given a limit on cost or area, select the
identify sets of nature reserves that maximize the representation of regional combination of sites that maximizes the
diversity, but, until recently, the methods have not dealt explicitly and directly representation of natural features (Box 2).
with the main goal of reserve networks, that of the long-term maintenance of
biodiversity. Furthermore, the successful application of current methods Advances in site-selection algorithms
requires reliable information about species distributions, which is not always Variants of these two problems have been tackled
available. Recent results show that data quality, as well as the choice of with different site-selection algorithms (Box 3). By
surrogates for biodiversity, could be critical for successful reserve design. the early 1980s, several authors1,3,4 independently
Because of these problems and a lack of communication between scientists developed iterative heuristic algorithms that attempt
and managers, the impact of computational site-selection tools in applied to solve the problem of combining sites into
conservation planning has been minimal. representative and efficient networks based on the
concept of COMPLEMENTARITY5,6. Subsequently, authors
Site-selection algorithms are computational methods began using tools from the field of OPERATIONS RESEARCH
Mar Cabeza*
Atte Moilanen that have been developed to identify a set of nature to contribute to the development of reserve design
Dept of Ecology and reserves containing as many species as possible, that algorithms. In the 1990s, discussions about the
Systematics, Division of is, methods maximizing the REPRESENTATION (see optimality properties of site-selection algorithms
Population Biology,
PO Box 17
Glossary) of species diversity1–3. For this purpose, two became a central theme in the literature, which
(Arkadiankatu 7), general representation problems have been concluded that heuristic algorithms do not guarantee
University of Helsinki, formulated. First, the minimum area problem: select an optimal minimum area solution, in contrast to
FIN-00014 Helsinki,
the sites that represent all NATURAL FEATURES a given exact algorithms (such as integer programming
Finland.
*e-mail: number of times with a minimum number of sites, techniques)7 (Box 3). Several studies2,3 have
cabeza@cc.helsinki.fi area or cost (Box 1). Second, the maximum coverage compared the optimality of different site-selection

http://tree.trends.com 0169–5347/01/$ – see front matter © 2001 Elsevier Science Ltd. All rights reserved. PII: S0169-5347(01)02125-5
 Review TRENDS in Ecology & Evolution Vol.16 No.5 May 2001
Box 1. Minimum area problem
Minimize number of sites, total area, or cost, to represent all natural
features (e.g. species)
A is an M sites x N natural features matrix whose elements aij are a measure
of the occurrence of the feature j in site i. Let Ii ∈{0,1} be an indicator variable
which has value 1 if site i is included in the selection and 0 otherwise. Let
each site have cost ci, and a desired representation level rj.. The
optimization problem is:
M
Minimize Eqn 1: ∑ ci Ii
i =1
M
subject to Eqn 2: ∑ aij Ii ≥ rj
i =1
for all j (each feature is represented at least rj times).
Particular problems of representation can be derived from this general casea:
(a) Minimize the number of sites needed to represent features:
ci = 1 all sites are of the same size or cost
aij = 0 or 1 if only the presence or absence of the feature is considered
rj = 1 gives the basic single representation problem
If rj ≥ 1 we have the multiple representation problem
(b) Minimize the total area needed to represent each feature at a level rj.
ci = areai the cost of a site is its area
aij = area covered by feature j in site i
rj = required representation level in area units
(c) Minimize total cost to represent features.
as (a) but the cost of a site is its precise real cost: ci = costI.
Reference
a Pressey, R.L. et al. (1997) Effectiveness of alternative heuristic algorithms for identifying
indicative minimum requirements for conservation reserves. Biol. Conserv. 80, 207–219
algorithms, giving mixed results. Although iterative
heuristic algorithms are often suboptimal, the
difference to the solution given by an exact algorithm
depends on the data set2,3.
The first systematic study comparing the solutions
found by different algorithms in relation to data-set
characteristics, such as rarity or nestedness of
features, size variation of the selection units and the
size of the data set, has only recently been conducted8.
The number of rare features (land types in this study)
in the data was an important factor in explaining
differences between iterative heuristics and integer
programming: the larger the number of rare features,
the more sites are needed and the closer are their
solutions. For instance, a high level of endemism
forces the initial inclusion of many sites into the
reserve network and, consequently, the remaining
optimization problem becomes computationally
simpler (and the results of different algorithms closer
to each other). Size variation among the sites also
appeared to reduce suboptimality when the goal was
to minimize the total reserve area rather than the
number of sites.
Although integer programming techniques should
be preferred for their guaranteed optimality, it has
been argued that they are not convenient, because
they may require unacceptably long computation
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243
Box 2. Maximal coverage problem
Maximize the representation of natural features
given a limit for the number of sites, cost or area
Let S be the set of sites included in the site
selection, N is the number of features. The object of
optimization is to:
N
Maximize Eqn 1:
∑ V j (S) [1]
j =1
[1]
∑ ci
subject to Eqn 2: [2]
≤R
i ∈S
[2] Vj (S) is the value of feature j in selection S. In the
simplest form, Vj (S) is 1 if the desired
representation level for feature j is achieved in
selection S and 0 if it is not. Vj (S) could also be a
measure of the predicted persistence of species j
on selection S. The cost of patch i, ci , can be defined
as 1 (if only the number of patches counts), as the
patch area, or as the real patch cost depending on
the objectives of optimization. R is the maximum
available resource, counted in the same units as ci.
(See Ref. a for detailed formulations.)
Reference
a Arthur, J.F. et al. (1997) Finding all optimal solutions to the
reserve site selection problem: formulation and
computational analysis. Environ. Ecol. Stat. 4, 153–165
times for large problems (hundreds or thousands of
sites or patches), and because they cannot solve
certain complex problem variants, including those
with nonlinear object functions2. Nevertheless, an
increasing fraction of conservation planning problems
can be solved using exact algorithms9, and there is
currently less emphasis on optimality, which is no
longer a problem in many cases. Integer
programming is preferred for both theoretical
exercises and when the size of the data set and the
conservation goal are appropriate9–11, whereas
iterative heuristics are employed when solutions are
required in seconds or minutes, and in conservation-
planning projects requiring close interaction with
decision makers4,12. Advanced heuristics, such as
stochastic global search methods (Box 3), appear to
represent a good compromise; these methods can deal
relatively quickly with complex problems and their
solutions are much less suboptimal than are those
given by the simplest iterative heuristics13. Also, in
recent reserve design literature, increasing attention
has been given to the use of multicriteria methods4,13.
Following algorithmic improvements and a great
increase in hardware speed, topics other than
algorithm efficiency and optimality are receiving
increased attention in the current reserve design
literature. These include the critical issue of how to
deal with long-term persistence of biodiversity, the
sensitivity of algorithms to data-set quality, the choice
of surrogate species for reserve design, and how to
244 Review TRENDS in Ecology & Evolution Vol.16 No.5 May 2001

Box 3. A phylogeny of site-selection algorithms

Ordinary single criterion optimization
minimizes (or equivalently maximizes)
one quantity. Single criterion optimization
methods that have been used for site-
selection problems can be categorized
into exact and inexact methods.
The branch-and-bound methoda of
integer programming is an exact method.
It considers all possible site selections
using implicit enumeration, and
guarantees to find the best solution(s).
Branch-and-bound methods require an
exponentially increasing number of
iterations as a function of problem size,
and therefore dataset size limits their
applicability.
Inexact heuristics (optimality is not
guaranteed and there is no estimate of the
quality of the solution) can be divided into
iterative heuristics and stochastic global
search methods. Iterative heuristics
repeatedly apply a set of rules, such as ‘add
the site that most complements the current
selection’, with no randomization used
except occasionally for breaking ties.
Iterative heuristics are very fast to
compute, but they can fail to find an
optimal solution even for simple
problemsb,c. Stochastic global search
techniques seek good solutions using
intelligent randomization. These
techniques are faster than integer
programming, but slower than iterative
heuristics. They cannot guarantee
optimality, but various difficult
optimization problems have been solved
successfully using stochastic global search
techniquesd.
Scoring methods are multicriteria
optimization methodse,f that consider
several objectives simultaneously (number
of sites, cost or persistence, etc.). Typically,
tradeoffs have to be made between different
objectives; for example, low cost might
prevent full biodiversity representation.
A multicriteria optimization problem
can be transformed into a single criterion
problem by taking a weighted sum of the
different objectives. Alternatively, a
constrained single criterion problem can
be defined by setting limits to all
objectives except one.
Note that all the methods that have been
used for the single criterion case can be
used to solve ‘subproblems’ in multicriteria
site-selection algorithms. However, in
multicriteria optimization, the critical phase
of optimization, is the determination of the
criteria and the weights given to them.
References
a Nemhauser, G.L. and Wolsey, L.A. (1988)
Integer and Combinatorial Optimization, John
Wiley & Sons
b Underhill, L.G. (1994) Optimal and suboptimal
reserve selection algorithms. Biol. Conserv. 70,
85–87
c Pressey, R.L. et al. (1996) Optimality in reserve
selection algorithms: when does it matter and
how much? Biol. Conserv. 76, 259–267
d Bäck, T. et al. (1997) Handbook of Evolutionary
Computation, Oxford University Press
e Steuer, R. (1986) Multiple Criteria
Optimization: Theory, Computation and
Application, John Wiley & Sons
f French, S. (1988) Decision Theory: An
Introduction to the Mathematics of Rationality,
Ellis Horwood

establish conservation-scheduling priorities given the
subset of selected sites.
Performance of existing reserves
The performance of existing reserves has often been
assessed by comparing them to the minimum number
of sites (or area) needed to represent all diversity
(efficiency sensu Pressey and Nicholls14), even though
minimal site number might not have been the original
goal of the reserve design. In practice, reserves have
frequently been selected in an ad hoc manner12,15,
without the use of explicit criteria. This explains the
observation that fewer sites are needed to represent all
diversity if the selection assumes no existing reserves
rather than beginning by including existing reserves15.
Although efficiency can be used as a measure of
the performance of an algorithm searching for the
minimum set (to compare solutions given by different
algorithms for the same data set), the success of a
reserve network depends on the initial conservation
objectives and, hence, success can and should be
measured in different ways. For example, Araújo16
and Rodrigues et al.10 have measured the success of
existing reserves by considering the gap between
current and complete representation, concluding
that, although the existing protected areas do not
sample all species, they do provide a better result
than does choosing areas at random
The assessment of the success of a reserve in
terms of persistence requires monitoring over time
to establish whether the conservation targets have
truly persisted10,16,17. A few recent studies have
demonstrated how efficient reserve networks
(efficient in representing numbers of species relative
to the cost) might not protect the original set of
features for many years, because species go locally
extinct and colonize sites regardless of whether the
site is included in the reserve network. Table 1
summarizes results from three studies that
analysed how well different reserve design
strategies would have maintained diversity over
time. The first two studies18,19 evaluated the
minimum area strategy in its ability to maintain
diversity. Both concluded that species turnover was
high, and that a large fraction of species could be lost
even in a short time (Table 1). The third example11
analysed the long-term performance of several
design strategies using the Common Birds Census
data set in Britain. This is the first study that
suggests ways of accounting for species turnover,
and it concludes that fewer species are lost if sites
are selected according to population density or local
abundance. The density strategy might avoid
selecting large sites with large populations, and
favour smaller sites with denser populations, and it
might, therefore, be a relatively efficient approach
(Table 1). However, more studies are needed to
establish whether focusing on good-quality (densely
populated) but possibly small sites would be a good
general strategy for reserve design. METAPOPULATION
THEORY suggests that many sites are needed for
species living as metapopulations in networks of

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 Review TRENDS in Ecology & Evolution Vol.16 No.5 May 2001 245

Table 1. Studies demonstrating how species tend to be lost from reserves in minimum set designsa

Study Criteria Species loss (%)b Reserve size Ref.

To minimize the number of sites that cover At least once 36.0 18 sites 18
all plant species from limestone pavements At least twice for species with >1 34.0 22 sites
(11 yr; 50 spp.; 77 sites)c population (78% of species)

For a combination of boreal lakes, find the Maximum number of plants 18.0 5 sites 19
minimum area that covers: Maximum restricted range diversity 18.0 5 sites
(63 yr; 32 spp.; 25 sites)

Minimize reserve network area to selectd All species at least once Min. Max. Mean Min. Max. Mean 11
(10 yr; 47 spp.; 56 sites) All species at least at the site where 4.0 12.0 8.0 400 750 556.25 (ha)
they are most abundant 0.0 4.0 2.7 1500 2250 1831.25 (ha)
All species at least at the site where 0.0 5.0 3.2 1000 1800 1312.50 (ha)
they have the highest density
aStudies analyzing the proportion of species that would have been successfully protected if the selection of the reserve had been done with the occupancy information at

some previous point.

bSpecies loss factors from Refs 18 and 19 are approximate values measured from graphs.

cTime intervals, number of features (species) and number of available sites.

dResults show minimum, maximum and mean data from eight analyses of data sets sampled ten years apart from a temporal data set of 20 years.

small patches, because the local extinction rates
would inevitably be high20.
Persistence of biodiversity in reserve networks
Researchers now agree that biodiversity persistence
should be considered in reserve-network
design4,12,13,17,21–23. So far, only a few studies have
actually done this. Some have focused only on local
(within site) persistence11,21,24,25, whereas others have
looked at regional (within reserve network)
persistence2,18,26. Spatial population dynamics,
however, has yet to be treated explicitly in site-
selection literature. Nevertheless, the general idea of
keeping sites close together has been influential in
spatial reserve design.
Local and regional persistence
Several approaches to deal with local persistence
have been suggested:
(1) Only selecting sites larger than a threshold size
when there is a known size limit for the presence of
species (e.g. the Schonewald-Cox index gives an
indication of the minimum area necessary for finding
particular mammal species)25.
(2) Basing site selection on abundance, either by
prioritizing sites with the largest populations11, or by
using population density as an indicator of site
quality11,21,27.
(3) Running population viability analysis for each
species24. As far as we know, this approach has not
been used in a multispecies site-selection exercise,
undoubtedly owing to the huge amount of high-
quality data that it requires.
If the focus of reserve design is on the regional
persistence (i.e. maintain each species within the
reserve network, even if local extinctions occur), a
preferred approach (assuming that the cost allows it)
is to include multiple representations of each natural
feature within the set of sites2,18,28. Another recent
study26 proposed a method that combines
probabilities of local persistence among sites directly,
until the required level of persistence probability for
each species is reached. The difficult part here is how
to obtain the probabilities of persistence. Araújo and
Williams29 used niche-based regression models to
estimate the local probabilities of occurrence of
species, which were then transformed into
probabilities of persistence with information on
threat and susceptibility of species to the threats.
Spatial reserve design
Economic considerations might require compact
reserves because the cost of management often scales
more closely with the length of reserve boundary than
with its area13. Moreover, a compact reserve increases
local persistence by reducing edge effects30 and, at the
same time, having individual sites close together
facilitates dispersal and recolonization of empty
habitats, which increases the probability of regional
persistence20,31. A few studies have considered the
clustering of sites by
• Including a rule that chooses sites close together
when breaking ties in iterative algorithms32,33.
• Minimizing a linear combination of reserve
network area and boundary length13.
Tight clustering of sites might not always be the
best way to increase persistence when REGIONAL
STOCHASTICITY is strong. Diseases, weather
catastrophes and forest fires, etc. can increase the
risk of simultaneous extinction of sites that are
located very close to each other34. One ad hoc solution
is to require multiple representations at sites
separated by a minimum distance while maximizing
reserve compactness13.
Despite these approaches to promoting
persistence, site-selection methods are essentially
based on static criteria, because they mostly use
information based on a single snapshot of species
distributions. Spatiotemporal dynamics are not
explicitly taken into account, even when species
distributions are known to change over time. Given
that many communities show high turnover18, the

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246 Review TRENDS in Ecology & Evolution Vol.16 No.5 May 2001
Glossary
Complementarity: Property of two (sets of) sites that occurs when some of the natural features
in a site differ from the features in another site. When (sets of) sites are highly complementary,
they contain (almost) nonoverlapping representation of natural features.
Irreplaceability: A measure of the likelihood that the site will be required as part of a reserve
network that satisfies a specific conservation goal. A site is highly irreplaceable when it includes
unique or rare natural features.
Metapopulation theory: In general, spatial population dynamics of a species living in a
spatially structured, possibly highly fragmented, landscape. A metapopulation may persist
regionally in a stochastic balance between extinctions of local populations and recolonizations
of empty habitat patches.
Natural features: Attributes (i.e. species, plant communities or landscape types, etc.) valued
for a conservation goal.
Operations research (OR): A mathematical discipline concerned with the construction,
mathematical analysis and solving of decision problems. OR seeks the determination of the best
(i.e. optimal) course of action under limited resources.
Regional stochasticity: Spatially correlated environmental stochasticity, caused for example
by weather phenomena. Regional stochasticity induces correlation into local population
dynamics and may cause simultaneous extinctions of closely spaced populations.
Representation: The extent to which required natural features occur within a (set of) sites.
Vulnerability: Risk of a site being transformed in a way that (some) natural features are lost
from the site.
sites selected by a static approach might not protect
as many species in the long term as the outcome of the
algorithm would have us believe. Awareness of this
problem has been growing and several
authors13,17,21,23 have called for the integration of
spatial population modelling in reserve network
design, which would require more interaction
between disciplines such as landscape ecology,
metapopulation biology and conservation biology.
In their recent study, Araújo and Williams29
attempt to deal with spatial dynamics in an implicit
way. They use spatial aggregation measures to
predict species occurrences, and they interpret spatial
correlations as a result of colonization processes. The
selected sites for the data set used in their study were
also clumped together, supporting the spatial design
approaches previously described. We are not aware,
however, of any studies incorporating spatial
dynamics explicitly in site-selection algorithms and
the problem remains a major challenge for future
reserve network designs. The critical part in such a
study is likely to be the acquisition of properly
parameterized species-specific models of spatial
dynamics for multiple species.
Data uncertainty
While site-selection algorithms continuously
improve, the quality of the biodiversity distribution
data needed for applying these methods remains poor.
In some cases, with incomplete biodiversity surveys,
data interpolation techniques have been used to
approximate species distributions3,35,36.
Risks associated with the use of incomplete data
sets in site-selection procedures have been recently
investigated28,37. In both studies, data sets were
systematically modified to assess the effects of
missing sites, missing taxa and missing records on
various variables. The first study37 concluded that
data deletions result in increased variation in
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selected networks and decreased efficiency because
more sites are needed to achieve the goal when the
level of deletion increases. The second study28
assessed the differences in selections in terms of the
percentage of the initial goal achieved (i.e. five
representations of each species from the complete
data set covered by the set selected with the reduced
data). In general, algorithms turned out to be
relatively robust to randomly distributed reductions
of records (up to 80–90% of the goal achieved) but
much less so when the missing data were
concentrated in particular sites or taxa.
The important conclusion is that inventory efforts
should be distributed as broadly as possible among
sites and taxa. In practice, the problem is often that
available data on species abundances and
distributions are biased towards preferred sites or
towards a few charismatic species. The use of
umbrella species as surrogates for poorly known
regional biota has been a common conservation
strategy38, comparable to the use of indicator taxa in
reserve design22. Andelman and Fagan39 evaluated
several umbrella species schemes and found that
none performed any better than did randomly
selected species from the database. Williams et al.40
reached the same conclusion in a similar study,
evaluating the use of flagship species to represent
other taxa. Williams et al.40 also found that areas
selected using flagships were no better for
representing biodiversity than was randomly
selecting the same number. Furthermore, sets of
complementary sites for different taxa often do not
overlap16,22,41,42, although results obtained in
Uganda43 do seem encouraging. In Uganda, different
taxa show similar biogeography, and therefore,
representative sets of sites for one taxon also
represent well other taxa. The conclusion from these
few studies is that the degree of overlap among taxa
varies depending on the geographical region and the
taxa being considered.
Another approach is to use habitat diversity as a
surrogate for species diversity. Although the approach
seems promising, results from recent studies40,44,
using species and environmental data for Europe44
and Africa40 are not encouraging. When applied to the
European data, the approach only performed well for
plants, with other taxa not showing a consistent
pattern of representation. The unrepresented species
mainly had restricted range sizes, were narrow
endemics, or were at the edges of their ranges. For the
African data, using ecoregions to select sites was a
better strategy to represent birds and mammals than
was using flagships, but still not as good as using data
for all birds and mammals.
Where to go from here
After two decades of research, site-selection
algorithms have improved to the point where they can
deal with a wide variety of design objectives. Despite
this, their impact in applied conservation planning
 Review TRENDS in Ecology & Evolution Vol.16 No.5 May 2001 247

Acknowledgements
We are grateful to
M. Araújo and I. Hanski for
discussion and comments
on this review. We also
appreciate the critical
comments of B. O’Hara,
H. Possingham and
R.I. Vane-Wright.
Financial support to MC
and AM was provided by
the Academy of Finland,
research projects #45125
and #71516, respectively.
has been minimal. The work done so far remains
mostly theoretical, with few exceptions2. Prendergast
et al.31 have discussed the gaps between theory and
practice in reserve design, and the lack of
communication between scientists and managers
appeared to be the main problem. Managers are not
aware of existing computational and analytical tools
and scientists remain unaware of many management
goals and constraints; neither is the optimal set of
sites selected by an algorithm generally readily
available45. The acquisition and management of sites
takes time. To cope with this, and to make decisions
about the scheduling of conservation action, the use of
IRREPLACEABILITY indices45,46 and VULNERABILITY indices
has been suggested12. This information can be
combined with other information (e.g. costs) in a
multicriteria decision analysis, which has recently
been suggested as a way of incorporating analytical
tools into a management framework4,12,47. Another
recent approach is the incorporation of site-selection
algorithms into decision-support systems to guide
negotiations between groups12.
Analytical reserve design tools are not very helpful
without high-quality data sets with which to apply
them. Ways of improving data quality include the
extension of surveys to avoid biases towards
particular taxa or sites28 and the use of statistical
tools to predict species distributions12,25. How to select
taxonomic groups used in reserve design remains
another problem12,22. Tests of taxonomic and
environmental surrogacy for species diversity are
generally discouraging, although some positive
results have been obtained43. The conclusion so far is
that good biodiversity surrogates are very difficult to
find.
Although data about the performance of existing
reserves are accumulating, more effort is needed in
monitoring reserves designed with site-selection
methods. All results from studies11,18,19 assessing the
long-term performance of reserves designed by site-
selection algorithms come from hypothetical
situations and not from monitoring of established
reserves. These studies have looked at how many
species would have persisted if the reserve network
had been designed using occupancy information at
some point in the past. Although all studies11,18,19
concluded that species turnover can be high, the
results concerning the performances of algorithms are
still likely to be optimistic. More species might have
disappeared if the remaining habitat not selected for
the reserve had deteriorated in quality. Such habitat is
likely to serve as a source of migrants that decreases
extinction probabilities in the reserve sites. With this
in mind, the explicit consideration of persistence and
spatiotemporal dynamics in reserve design remains a
major future challenge for the discipline.

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Linking plants to rocks: ectomycorrhizal

fungi mobilize nutrients from minerals
Renske Landeweert, Ellis Hoffland, Roger D. Finlay, Thom W. Kuyper and Nico van Breemen

Plant nutrients, with the exception of nitrogen, are ultimately derived from accelerate weathering of minerals by excreting
weathering of primary minerals. Traditional theories about the role of organic acids, phenolic compounds, protons and
ectomycorrhizal fungi in plant nutrition have emphasized quantitative effects SIDEROPHORES3,4. Such plant-induced mineral
on uptake and transport of dissolved nutrients. Qualitative effects of the weathering must have facilitated the subsequent
symbiosis on the ability of plants to access organic nitrogen and phosphorus spread of land plants by increasing the availability of
sources have also become increasingly apparent. Recent research suggests essential plant nutrients and by producing SECONDARY
that ectomycorrhizal fungi mobilize other essential plant nutrients directly MINERALS, such as clays and iron (Fe) and aluminium
from minerals through excretion of organic acids. This enables ectomycorrhizal (Al) oxides, providing soil material for anchorage and
plants to utilize essential nutrients from insoluble mineral sources and affects water holding. Continuing today, essential plant
nutrient cycling in forest systems. nutrients enter ecosystems through biogeochemical
weathering of primary minerals. To understand soil
Physical and chemical WEATHERING (see Glossary) of ecological processes, knowledge of the potential
Renske Landeweert* the primary rock of the earth results in a release of influence of plant roots and their associated
Thom W. Kuyper dissolved mineral elements and residues into the microbiota on weathering processes is essential.
Sub-dept of Soil Quality,
Wageningen University,
biological environment. The rise of vascular land
Box 8005, NL-6700 EC plants about 400 million years ago (Mya) presumably Organic acids as weathering agents
Wageningen, led to vastly increased mineral weathering1,2. Some of Soluble organic acids affecting mineral weathering in
The Netherlands. the photosynthetic products formed in plant leaves soils originate from various sources. Medium to high
*e-mail:
Renske.landeweert@
end up as organic acids that are exuded by plant roots molecular weight organic acids, such as humic
bb.benp.wag-ur.nl into the surrounding soil. Together with CO2, which is substances, are less effective in promoting mineral
Ellis Hoffland
pumped into the soil via root respiration and dissolution than are low molecular weight (LMW)
Nico van Breemen heterotrophic respiration of soil microorganisms, organic acids produced by plant roots and soil
Laboratory of Soil Science these organic acids greatly enhance the dissolution of microorganisms5. Although constituting only a minor
and Geology,
primary silicate minerals (Box 1), thereby mobilizing fraction of the total organic acids in the soil solution,
Wageningen University,
Box 37, NL-6700 AA essential LITHOPHILIC plant nutrients. Silicate LMW organic acids are generally considered to be the
Wageningen, minerals such as FELDSPARS, MICAS, HORNBLENDE and most important biological weathering agents in soils,
The Netherlands. pyroxene provide calcium (Ca), magnesium (Mg) and owing to their acidifying and complexing capacities5,6.
Roger D. Finlay potassium (K); APATITE is the main PRIMARY MINERAL Depending on the number and configuration of
Dept of Forest Mycology source of phosphorus (P). carboxylic and phenolic groups and the related acid
and Pathology, Swedish
The carbon-rich root exudates also support large strength, organic acids provide H+ for protonation of
University of Agricultural
Sciences, Box 7026, SE- communities of root-associated microorganisms the mineral surface and chelate cations7 (Box 1).
750 07 Uppsala, Sweden. (rhizosphere bacteria and fungi) that further Concentrations of LMW organic acids in the bulk soil

http://tree.trends.com 0169–5347/01/$ – see front matter © 2001 Elsevier Science Ltd. All rights reserved.PII: S0169-5347(01)02124-3