866 Ecological Processes | Denitrification

Denitrification
U Skiba, Centre for Hydrology and Ecology, Penicuik, UK
ª 2008 Elsevier B.V. All rights reserved.

Denitrification Further Reading

Denitrification aerobic heterotrophs in soil than they are as denitrifiers.

However, the ability to respire using O2 or NO3 enables
Denitrification is the microbial process of reducing nitrate denitrifiers to survive and proliferate in a wide range of
and nitrite to gaseous forms of nitrogen, principally oxic, micro-oxic, and anoxic environments and adapt
nitrous oxide (N2O) and nitrogen (N2). A large range of quickly in rapidly changing environments.
microorganims can denitrify. Denitrification is a response The final product of denitrification is not always N2, but
to changes in the oxygen (O2) concentration of their could be N2O and sometimes even NO, depending on the
immediate environment. Only when O2 is limited will organism involved and on the environmental conditions.
denitrifiers switch from aerobic respiration to anaerobic Equally, NO3 does not need to be the terminal electron
respiration, using nitrate (NO3) as electron acceptor. acceptor; instead, other intermediates may be used. For
example, the bacteria Alcaligenes faecalis, Pseudomonas stutzeri,
Paracoccus denitrificans, and Pseudomonas perfectomarinus were
Process of Denitrification shown to grow in the laboratory using N2O as the electron
acceptor.
The classic definition of denitrification is the stepwise
reduction of nitrate (NO3– ) to dinitrogen gas (N2)
Nitrifier denitrification
nitrate ! nitrite ! nitric oxide ! nitrous oxide Nitrifier denitrification is a pathway carried out by the
! dinitrogen gas group of nitrifiers that perform the first step of nitrifica-
tion, the ammonium oxidizers, which oxidize ammonium
NO3– ! NO2– ! NO ! N2 O ! N2 (NHþ –
4 ) to hydroxylamine (NH2OH) and then NO2 .
–
Under O2 stress, these organisms then reduce NO2 to
The reductions are catalyzed by the enzymes nitrate
gaseous forms of N (NO, N2O, N2):
reductase, nitrite reductase, nitric oxide reductase, and
nitrous oxide reductase. The redox balance of denitrifica- NHþ –
4 ! NH2 OH ! NO2 ! NO ! N2 O ! N2
tion is
The main nitrifying species are autotrophic bacteria; they
2NO3– þ 10e – þ 12Hþ ! N2 þ 6H2 O use CO2 as a carbon source. Nitrifier denitrifiers can use O2
for two separate functions, as a substrate in the oxidation of
NHþ 4 and subsequently as a terminal electron acceptor in
Microorganisms Involved in Denitrification respiration. It has been suggested that this process can con-
Pathways tribute significantly to the loss of NHþ
4 as NO and N2O from
Heterotrophic denitrification soil; however, it is difficult to really distinguish between
A large range of heterotrophic microorganisms can autotrophic nitrifier denitrification and heterotrophic deni-
denitrify. Heterotrophs are organisms that require trification. The nitrite reductase enzyme of nitrifers, and
organic carbon supply for growth. Denitrifiers are very classic denitrifiers are biochemically and genetically similar.
successful and are present in high numbers in both For example, similar spectroscopic characteristics, inhibition
terrestrial and marine ecosystems in all climate zones profiles, and reaction products were demonstrated for the
of our globe. They include bacillus, paracoccus, pseudo- copper-containing enzyme of Nitrosomonas europaea. Genetic
monas, propionibacterium, and thiobacillus species. Most similarities were observed between the nirK gene of a classic
denitrifiers are facultative aerobes, which means that denitrifier and Nitrosomonas marina and other closely related
they gain energy using O2 as the electron acceptor ammonium oxidizers.
(aerobic respiration), but are able to switch to anaerobic
respiration and use NO3 as an electron acceptor when
O2 becomes depleted. The enzymes required for deni- Chemodenitrification
trification are activated by lowering the oxygen partial Chemodenitrification is the chemical, nonenzymatic
pressure. Denitrifiers are generally more competitive as decomposition of intermediates from the oxidation
 Ecological Processes | Denitrification 867

of ammonium to nitrite and nitrate (nitrification). This can be a real problem, but direct measurements have been
reaction usually occurs at low pH and may be facilitated achieved by several scientific groups who replaced the
by the presence of iron or amines and high levels of atmosphere of soil columns with helium and then directly
organic matter. The most important chemodenitrification measured the N2 resulting from denitrification.
reaction is the decomposition of HNO2 to form NO. The traditional methods to measure total denitrifica-
tion are by the acetylene block technique or by adding
3HNO2 ! 2NO þ HNO3 þ H2 O 15
N as a tracer. Acetylene at a partial pressure of 10 kPa
Soil pH is a controlling factor in chemodenitrification blocks the final step of denitrification, so that the total
and above pH 5, this process is insignificant. product of denitrification is N2O. The main problem of
this method is that it is not very easy to administer the
inhibitor evenly throughout soils. The most active deni-
Measurements of Denitrification trification zones in a soil are likely to be those into which
diffusion is limited and have effectively managed to
The main products of heterotrophic denitrification are exclude O2.
N2O and N2 and of nitrifier denitrification, they are NO In tracer experiments, nitrogen substrates are labeled
and N2O. Emissions of NO and N2O are measured rela- with 15N and added to soils or waters. The appearance of
tively easily. For terrestrial systems a small area of the soil this label in the denitrification products N2O and N2 can
surface is enclosed by a chamber (see Figure 1), or in the be measured by isotope-ratio mass spectrometry.
laboratory a small amount of soil is incubated in an air- There are several methods that are used to distinguish
tight vessel, in order to allow the gases emitted to increase between N2O production by autotrophic nitrifiers or het-
to measurable concentrations. N2O is a very stable gas and erotrophic denitrifiers. (1) Acetylene at a partial pressure
samples from the enclosures can be stored in syringes or of 10 Pa inhibits N2O production during chemoauto-
vials until analysis by gas chromatography, using an trophic nitrification, whereas a much higher partial
electron-capture detector. NO is a highly reactive gas, pressure of 10 kPa is required to inhibit the reduction of
with a half-life of less than 1 min and therefore can only N2O to N2 in heterotrophic denitrifiers. (2) The contri-
be measured directly from the enclosed chamber by bution of nitrifiers and denitrifiers to N2O can be studied
chemiluminescence. In aquatic systems, the concentration by differential labeling NH4 and NO3 with 15N and
of N2O/NO in the water columns or in the headspace monitoring the occurrence and disappearance of the 15N
above the water column is analyzed. label in the NH4, NO3, and N2O pools. (3) Inhibitors
The measurement of the final denitrification product which specifically inhibit autotrophic nitrification (nitra-
N2 is not so straightforward. Direct measurements can pyrin), denitrification (O2), or heterotrophic nitrification
only be made in an N2-free environment. This is difficult, (cycloheximide) have been used to distinguish between
as more than 80% of our atmosphere is N2. Contamination these processes. (4) The use of natural abundance isotope
ratios is a new approach, still in development. The frac-
tionation of isotopic ratios 14/15N and 16/18O is different
during nitrification and denitrification.

Requirements for Denitrification to Occur

The basic requirements for denitrification to occur in any
marine, freshwater, or soil environment, regardless of
climate are reduced O2 concentrations, a nitrogen source,
and available carbon source.
Nitrate, the substrate for heterotrophic denitrification
stimulates denitrification and large rates of denitrification
are observed in agricultural soils, wastewater treatment
farms, N polluted streams, estuaries, and forests.
Figure 1 This is a typical chamber to measure the fluxes of However, even when the supply of NO3 is not limiting,
nitrous oxide from soil. The lid is closed for periods of 30–60 min, denitrification will only take place when the environmen-
gas samples are collected from the chamber at the beginning tal conditions activate and stimulate the denitrifying
and end of chamber closure through the sample port using a population to switch from aerobic respiration to anaerobic
syringe. Samples are stored in vials or gas sample bags and are respiration. If a soil is too dry or an aquatic system too
analyzed in the laboratory for nitrous oxide mostly by gas
chromatography. Usually, information of the soil temperature, aerated, microorganisms will not switch to denitrification.
soil moisture, and soil mineral NO3 and NH4 are collected at the Furthermore, a lack in labile organic carbon or increased
same time to provide a picture of the controlling variables. competition between heterotrophic bacteria for labile
868 Ecological Processes | Denitrification

carbon or competition with organisms that dissimilate (a) 300

Manure
nitrate to ammonium, will reduce denitrification.
250 NH4NO3

Nitrous oxide emission

Control of denitrification in terrestrial systems 200

(µg N ha–1 d–1)

Nitrogen
150
Agricultural soils receive regular N applications of mineral
N or organic fertilizers (farmyard manure, sewage sludge) 100
and in some systems by intercropping, or crop rotations,
50
with leguminous N fixing plants. The application of
N fertilizers stimulates nitrification and denitrification 0
activity. Under optimal soil conditions, when soil moisture

33
40
11
12
16
19
23
26
–8
1
3
4
5
6
7
9
–2
–50
and soluble carbon content are not limiting, the response to Days after spring fertilizer application in 2002
mineral N fertilizer application is very fast, resulting in
peaks of NO, N2O, or N2 emissions within hours of appli- (b) 160
Manure
cation. These emission peaks can be 100 times larger than 140
NH4NO3
background emissions, but are short-lived and usually last

Nitrous oxide emission

120
for 1–3 weeks, before returning to near background emis-

(µg N ha–1 d–1)

100
sions. The magnitude and shape of the peak and the
dominant N gas emitted are controlled by the aeration of 80
the soil and the availability of organic carbon. A mineral 60
fertilizer provides a readily available N source for denitri- 40
fiers to utilize immediately, resulting in a sharp response, as 20
shown in Figure 2 in 2002. In 2003, there was very little
0
rainfall during fertilization. The N2O emission peak was –8
0
1
2
7
10
13
15
17
21
27
29
35
38
44
delayed until rainfall, but was smaller, because of uptake of –20
Days after spring fertilizer application in 2003
fertilizer N by plants and microbial biomass during the
period unsuitable for denitrification. Organic N fertilizers Figure 2 The influence of mineral N fertilizer, manure, and
provide not only a source of N, but also a source of carbon; rainfall on the emissions of N2O from intensively managed
grassland. Measurements were made using static chambers
the bulk of it needs to be mineralized before it is available
(Figure 1) before and at intervals after fertilizer application. The
for denitrification. This is the reason for the delayed, but differences in the shape and magnitude of the N2O peaks can be
much broader N2O emission peak due to denitrification in explained by differences in rainfall pattern. (a) In 2002, rainfall
Figure 2. distribution before, during, and after fertilizer application ensured
In other soils, nitrogen is supplied by mineralization of optimal conditions for N2O emission. (b) In 2003, fertilizer was
applied during a 3 week dry period, and rainfall only started
organic material or deposition of atmospheric nitrogen.
10 days after fertilizer application. The much broader peaks
Overall denitrification rates are much lower than that for associated with the manure treatment is a result of the organic
agricultural soils. carbon source supplied in manure, stimulating denitrification.
Modified from Jones SK, Rees RM, Skiba UM, and Ball BC (2007)
Oxygen Influence of organic and mineral N fertilizer on N2O fluxes from a
temperate grassland Agriculture, Ecosystems and Environment
Oxygen is the primary controller of denitrification.
121(1-2): 74–83.
Denitrification only operates when the soil atmosphere
O2 concentration ranges from 4% to 17%. The main
reactions stop at N2O whereas at high WFPS (>70%)
variables that control the soil O2 concentration are rain-
denitrification to N2 may dominate, provided the C and
fall, drainage, soil compaction, soil texture, and soil
N supplies are not limiting.
respiration. Rainfall, lack of drainage, and compaction
all reduce the O2 concentration in the soil and stimulate
denitrification. Fine-textured soils, with high clay con- Carbon
tent, are more readily depleted of oxygen than a coarse- The primary requirement for heterotrophic denitrifiers is
textured soil with high sand content. The lack of O2 of an easily available carbon source, which will fulfill two
a soil is often expressed in terms of its water-filled pore roles: (1) simple organic carbon compounds provide a
space (WFPS). This variable is calculated from the bulk carbon source for growth of heterotrophic denitrifiers
density and particle density of the soil and the water and (2) in an aerobic environment labile organic carbon
content. Denitrification operates when the WFPS is stimulates the respiration of O2 which in turn will reduce
greater than 55%. The ratio of the products of denitrifi- the O2 concentration surrounding the respiring bacteria.
cation N2O/N2 decreases as the WFPS increases. At the The denitrifiers among them eventually are under pres-
lower WFPS rates (<70%) most of the denitrification sure to switch from aerobic to anaerobic respiration using
 Ecological Processes | Denitrification 869

NO3. Many studies have shown an increase in denitrifica- leading to the emission of N2O from the water.
tion rate when a labile carbon source was added to soils in Denitrification occurs predominately in the top few mm
the laboratory or in the field. The importance of labile of the sediments and rates of denitrification depend on the
organic matter was shown in studies on intact soil cores, concentrations of organic C and N. Largest denitrification
and subsequently on dissecting these into smaller and rates can be expected from the most polluted rivers and
smaller subcores. Between 25% and 85% of the denitri- lakes and estuaries followed by coastal shelves and lowest
fication activity could be attributed to a single decaying rates from the open oceans. The main sources of NO3 are
leaf or worm. This study and many other field studies drainage and runoff from agriculturally managed fields
have shown that denitrification is a very opportunistic and effluents from sewage farms and industries. For rivers,
behavior, leading to spatial and temporal hotspots of the denitrification rates were shown to be inversely related to
production of N2O and N2. channel size, which would regulate the residence time of
Generally, the rhizosphere, the soil layer rich in root the NO3.
exudates and therefore high microbial activity, provides Estuaries have a relatively high organic carbon content
an ideal environment of high C and low O2 concentra- compared to many other aquatic systems, due to high
tions for denitrification to take place. The dense root mat rates of sedimentation and upwelling. They can denitrify
of grasslands is particularly active and denitrification rates between 20% and 50% (and more in isolated cases) of the
are larger than those in arable soils or forests. NO3 added by the rivers and thus stop the export of
Mineralization of complex carbon compounds in soil terrestrial N pollution to open oceans. Denitrification
increase the pool of simple carbon compounds. The most appears to be independent of salinity over the range of
well-studied processes that stimulate the mineralization 1–13 parts per trillion (ppt), but directly dependent on
of organic carbon are plowing, freeze–thaw events, and nitrate concentration.
wetting of very dry soil (pulsing effect). Plowing aerates In the interior of oceans, away from the euphotic surface
the soil and thereby stimulates mineralization of organic layer and coastal shelves, nitrification and denitrification
carbon compounds in parts of the soil previously not rates are primarily dependent on the mineralization of
receiving the O2 concentrations required for mineraliza- nitrogen-rich organic matter originating from N2 fixing
tion to take place. Increased mineralization stimulated by organisms. The demand for anaerobic conditions is
freeze–thaw events or the pulsing effects are governed by required in the same way as for terrestrial systems.
the same principle. Extreme cold and extreme dry condi- Denitrification rates in benthic sediments are most sensitive
tions kill a large part of the microbial community and to concentrations of labile organic carbon at the sediment–
reduce the activity of the survivors. When conditions water interface and have been shown to increase with
become more tolerable, due to a thaw or rainfall, the nitrate concentrations. The major areas of low O2 are the
remaining microbial population become activated and thermocline of the Arabian Sea and the eastern tropical
feed off the nutrients supplied by the dead microbial South and North Pacific. Many regions of the world’s
biomass and any nutrients deposited in the soil during oceans do not denitrify or fix nitrogen.
periods of drought or freeze. These three events are Measurements of the isotopic ratios 14/15N and 16/18O of
usually accompanied by increased rates in soil respiration N2O originating from surface and from deep waters in the
and NO and N2O emissions. Increased NO, N2O, and subtropical North Pacific Ocean have revealed large differ-
CO2 emissions are observed for a few days only and fade ences. Surface water N2O was depleted in 15N and 18O
away when the supply of nutrient has diminished. compared to atmospheric and deep water N2O. This
Subsequent pulsing, freeze–thaw cycles produce smaller implies that nitrification, rather than denitrification, is prob-
emission peaks, due to less substrate stored, compared to ably the main source of surface water N2O in open oceans.
the first event. The dominant N gas emitted depends on
the soil properties and meteorological conditions, with Control of denitrification by pH and temperature
low aeration stimulating N2O/N2 emission, while high pH
aeration stimulating NO emission. Emissions can be very Until recently, the general view was that in soils, optimal
high and these very sporadic events can significantly denitrification rates occur at pH 7–8, that the ratio of deni-
contribute to the overall annual NO or N2O flux. trification products N2O/N2 decreases as the pH is raised,
and that N2O produced in acid soils is more likely a product
Control of denitrification in aquatic systems of heterotrophic nitrification rather than denitrification.
In aquatic environments, the requirements for denitrifica- However, new studies have shown that microorganisms can
tion are the same as in terrestrial systems: a nitrogen and denitrify in acid soils, can adapt to acid soil conditions, and
carbon source and low O2 concentrations. The major end that microbial pH optima are similar to the pH of soils. Also,
product of denitrification in aquatic systems is N2, but in aquatic systems, a high denitrification potential was
surface layers of the water columns are also often observed for acid mine sediments from several affected
enriched in N2O in relation to the atmosphere, thus streams ranging from pH 2.6 to 4.9 in Colorado, US.
870 Ecological Processes | Denitrification

In contrast to the above, various reports have demon- estuaries with good light penetration, the diurnal pattern
strated that NO3 concentrations can be reduced most of benthic microalgal production and associated rise in O2
efficiently at higher pH in effluent treatment works. In concentration at the sediment surface, during the day
one study, denitrification of synthetic wastewater with a inhibits denitrification. Consequently, larger denitrifica-
high NO3 concentration was inhibited at pH values below tion rates can be observed at night. For rivers, however,
7, but complete denitrification occurred from pH 7.5 to 9.0. the opposite was observed. Denitrification rates were
The process of denitrification increases the pH of its higher during the day than night, in spite of maximum
surrounding environment, which can be beneficial for O2 concentrations during the day. It has been suggested
some aquatic systems. The buildup of high NO3 concen- that diurnal denitrification patterns are controlled by the
trations in a closed seawater aquarium was minimized by nitrification cycling driven by diurnal variations in water
installing a biological denitrification system. In addition, pH and temperature.
this system offers additional benefits by increasing the pH
and alkalinity of seawater and providing a reducing envir-
onment to balance the oxidizing disinfection environment
Models of Denitrification
in the aquarium.
The importance of denitrification as a source of atmo-
spheric N2O, a greenhouse gas, and also an effective
Temperature
removal process of harmful NO3 in aquatic systems has
All microbial activity, including denitrification, is influ-
stimulated many researchers into modeling denitrifica-
enced by changes in temperature. Activity is very slow
tion rates and its products. Modeling denitrification is
below 5  C, increases linearly to a maximum around
complicated, as two microbial processes with very con-
25–30  C and thereafter decreases to a minimum at
trasting requirements (nitrification and denitrification)
around 65  C, where growth stops due to enzyme dena-
have to be considered. The denitrifying population is
turation. There are exceptions and some bacteria have
always present and respiring, but will only denitrify
evolved to grow in extremely hot (>100  C) or cold
under very specific conditions. Denitrification is therefore
(–12  C) conditions. Temperature optima in a particular
a very transient process both in space and time.
soil appear to be the same for a range of microbial
For terrestrial systems, simple models calculate deni-
processes, including nitrification and denitrification, but
trification from few essential parameters, for example,
temperature optima differ for different soils. The tem-
NO3 and carbon content, soil texture, and drainage.
perature optima do not appear to be linked to climate.
More complex models describe the denitrification process
Strong seasonal and diurnal variations in N2O and NO
in detail, for example, the DNDC model (denitrification
fluxes are observed in all marine and terrestrial systems,
decomposition model).
but only when carbon and nitrogen supply and anaerobi-
In aquatic systems, denitrification has been calculated
city is not limiting. Under nonlimiting conditions, the
from diffusion of substrates and products to and from the
temperature coefficient of denitrification, Q 10, is mostly
active sites for estuaries. For rivers, N removal was cal-
reported around 2–4. Frequently, the temperature
culated from the physical properties of the stream,
response of denitrification and other microbial reactions
especially channel size. For marine sediments, a more
is expressed as an activation energy, which can be calcu-
complex model has been developed that couples the
lated from the plot of the natural logarithm of the flux
biogeochemical cycling of sedimentary carbon, nitrogen,
against the inverse absolute temperature. Usually, activa-
and oxygen with that in the water column models.
tion energies range from 50 to 100 kJ mol1, and again are
similar for a range of microbial processes.
See also: Ammonification; Biological Nitrogen Fixation;
Nitrification.
Diurnal patterns
Diurnal patterns of N2O and NO emissions from soils are
linked to diurnal oscillations of the soil temperature. Soil Further Reading
temperature diurnal oscillations dampen with increasing
Casciotti K and Ward BB (2001) Dissimilatory nitrite reductase genes
soil depth. Comparison of diurnal patterns of temperature from autotrophic ammonia-oxidizing bacteria. Applied Environmental
at a range of depth with diurnal patterns of N2O or NO Microbiology 67(5): 2213–2221.
emissions may be used to locate the depth of maximum Davidson EA (1991) Fluxes of nitrous oxide and nitric oxide from
terrestrial ecosystems. In: Rogers JE and Whitman WB (eds.)
denitrification or nitrification activity. In a well-drained Microbial Production and Consumption of Greenhouse Gases:
soil, denitrification occurs at deeper soil layers and nitri- Methane, Nitrogen Oxides and Halomethanes, pp. 219–234.
fication in the upper soil layers. Washington, DC: ASM Press.
Gruber L and Sarmiento JL (2005) Global patterns of marine nitrogen
In aquatic systems, very different diurnal denitrifica- fixation and denitrification. Global Biogeochemical Cycles
tion patterns are reported. In shallow streams, lakes, or 1: 235–266.
 Ecosystems | Desert Streams 871

Hofstra N and Bouwman AF (2005) Denitrification in agricultural soils: Middelburg JJ, Soetaert K, Herman PMJ, and Heip CHR (1996)
Summarizing published data and estimating global annual rates. Denitrification in marine sediments: A model study. Global
Nutrient Cycling in Agroecosystems 72(3): 267–278. Biogeochemical Cycles 10: 661–674.
Jones SK, Rees RM, Skiba UM, and Ball BC (2007) Influence of organic Seitzinger SP, Kroeze C, and Styles RV (2000) Global distribution of N2O
and mineral N fertilizer on N2O fluxes from a temperate grassland. emissions from aquatic systems: natural emissions and
Agriculture, Ecosystems and Environment 121: 74–83. anthropogenic effects. Chemosphere – Global Change Science
Klemedtsson L, Svensson BH, and Rosswall T (1988) A method of selective 2: 267–279.
inhibition to distinguish between nitrification and denitrification as Simek M, Jisova L, and Hopkins DW (2002) What is the so-called
sources of nitrous oxide in soil. Biology and Fertility of Soils 6: 112–119. optimum pH for denitrification in soil? Soil Biology and Biochemistry
Kuenen JG and Robertson LA (1988) Ecology of nitrification and 34: 1227–1234.
denitrification. The Nitrogen and Sulphur Cycles, pp.161–218, 42nd Skiba U and Smith KA (2000) The control of nitrous oxide
Symposium of The Society for General Microbiology, University of emissions from agricultural and natural soils. Chemosphere
Southhampton, Cambridge University Press. 2: 379–386.
Laursen AE and Seitzinger SP (2004) Diurnal patterns of denitrification, Wrage N, Velthof GL, van Beusichem ML, and Oenema O (2001) Role of
oxygen consumption and nitrous oxide production in rivers nitrifier denitrification in the production of nitrous oxide. Soil Biology
measured at the whole-reach scale. Freshwater Biology and Biochemistry 33: 1723–1732.
49(11): 1448–1458. Zehr JP and Ward BB (2002) Nitrogen cycling in the ocean: New
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Desert Streams
T K Harms, R A Sponseller, and N B Grimm, Arizona State University, Tempe, AZ, USA
ª 2008 Elsevier B.V. All rights reserved.

Distribution and Physical Template Nutrient Dynamics

Temporal Dynamics Human Modifications
Biota Further Reading
Energetics

Distribution and Physical Template precipitation regimes as well as in geomorphology. Thus,

the hydrogeomorphic templates and resulting ecological
Desert streams occupy arid and semiarid regions defined characteristics of desert streams exhibit a great diversity
by low annual precipitation. Semiarid and arid climate of patterns. Despite this extensive distribution of desert
zones are found on all continents and include both hot streams, the vast majority of ecological studies of desert
and cold deserts. Although the range of temperatures streams have occurred in the southwestern United States,
varies across desert regions, summer temperatures may Australia, and Antarctica. Our discussion thus draws from
exceed 40  C in hot deserts. Annual precipitation ranges results of studies in these ecosystems. Future studies of
from <100 to 300 mm yr1 and combined with high tem- desert stream ecosystems in other regions are likely to add
perature can result in high rates of evapotranspiration. new dimensions to the state of our understanding pre-
Higher precipitation in the mountains (up to 1000 mm sented here.
yr1) can feed streamflow in the low deserts, often sup- Desert stream hydrographs are punctuated by events
porting perennial flows in large basins. Arid and semiarid when discharge may exceed baseflow by several orders of
regions are characterized by distinct seasons defined by magnitude. Precipitation falling on the catchment rapidly
precipitation and/or snowmelt and the amount of preci- reaches the stream and stream discharge rapidly dissipates
pitation that falls during these seasons shows high following floods. Infiltration of desert soils is minimal and,
interannual variability. This results in extreme seasonal at the scale of whole basins, much of the water that is not
and interannual variation in stream discharge. Indeed, returned to the atmosphere by evapotranspiration reaches
streams in some desert regions flow in response to rain streams via overland flow during storms or via infiltration
events that occur only once in several years or even less. of permeable low-order channel sediments followed by
Arid and semiarid lands account for over one-third of subsurface flow. Resulting flash floods scour the
global lands, making desert streams prominent among streambed, resulting in downstream export of sediments
aquatic ecosystems. The large geographic area covered and aquatic organisms and creating a wide channel. Large
by deserts results in a wide variation in temperature and floods also deposit alluvial materials in riparian zones and