Hydrobiologia 482: 3139, 2002.

2002 Kluwer Academic Publishers. Printed in the Netherlands.

31

Ammonium retention and whole-stream metabolism in cave streams

Kevin S. Simon1 & Ernest F. Benfield
Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061, U.S.A.
1 Present address: Department of Zoology, University of Otago, Dunedin, New Zealand
E-mail: kevin.simon@stonebow.otago.ac.nz
Received 23 October 2001; in revised form 15 May 2002; accepted 27 May 2002

Key words: cave, stream, nitrogen, respiration, nutrient uptake

Abstract
We used 4 short-term ammonium releases and 1 whole-stream metabolism estimate to examine nutrient and carbon
dynamics in cave streams. Ammonium uptake lengths (34157 m) in the cave streams fell within the range reported
for surface streams. However, uptake lengths in the cave streams were longer than would be expected in surface
streams of similar size. Ammonium mass transfer coefficients and uptake rates were much lower in the cave
streams than have been measured in surface streams. Differences in ammonium uptake parameters among cave
streams were due to both abiotic and biotic factors. Metabolism in the cave stream was relatively low (32.9 g C
m2 y1 ), but fell within the range of values reported for surface streams. Benthic organic carbon (BOC) standing
stock was low and turned over faster in the cave stream than in most surface streams. Our results suggest cave
streams are carbon, but not nutrient, limited and that standard methods and models for nutrient and carbon cycling
in surface streams are applicable to cave streams.

Introduction
Much biological research has focused on the evolutionary, population, and community ecology of caves
(e.g. Culver et al., 1994; Gibert et al., 1994). In contrast, ecosystem processes, particularly carbon flux
and nutrient cycling, are almost unstudied in caves.
This is surprising considering caves have long been regarded as energy limited because primary production
is absent and organic matter sources are usually scarce
in caves. If caves are truly energy limited, carbon flow
and nutrient cycling should play important roles in the
ecology of cave ecosystems. Organic carbon entering
karst aquifers from the surface is oxidized (Albric
& Lepillier, 1998); however, little is known about
the biological importance and fate of organic matter
and nutrients after they enter the subsurface. This is
particularly important in karst because most surface
water is diverted below ground before eventually being removed through wells for human consumption
or emerging in springs or hyporheic zones of streams
(White, 1988). Nutrient and organic pollution of caves

is common and directly affects cave communities and

water quality in karst landscapes (Simon & Buikema,
1994; Boyer & Pasquarell, 1996). Factors affecting
the dynamics of carbon and nutrients in caves need
to be studied to understand what fuels and controls
cave communities and to understand nutrient and carbon delivery to surface systems that are connected to
groundwater (Gibert et al., 1990).
Carbon flow and nutrient cycling in surface
streams are well studied (e.g. Newbold, 1992; Webster & Meyer, 1997a) and the methods and models
developed for surface streams should be applicable to
cave streams. Nutrients in streams cycle between abiotic and biotic components while continuously or periodically traveling downstream. This process, called
spiraling (Webster & Patten, 1979; Newbold, 1992),
can be quantified as spiraling length, the distance
traveled during a cycle between abiotic and biotic
form. Most of the spiraling length consists of the distance traveled in abiotic form, called uptake length
(Sw ). Uptake length indicates the nutrient retentiveness (i.e. nutrient retentive streams have short uptake

32
Table 1. Size, flow, background nutrient concentration and NH4 uptake parameters (Sw , vf and U ) of the cave streams. SRP = soluble
reactive phosphorus. Sw = uptake length. vf = mass transfer coefficient. U = uptake rate. Uptake lengths with same superscript letter are not
significantly different (ANCOVA, p > 0.05). Values in parentheses are 1 SE for nutrient concentrations and 95% CI for Sw
Parameter
Mean width (m)
Mean depth (m)
Velocity (m/s)
Discharge (L/s)
NH4 + N (g/L)
NO3 N (mg/L)
SRP (g/L)
Sw (m)
vf (104 m/s)
U (g N m2 s1 )

S3

JC

LE

SRS

1.0
0.030
0.050
1.38
0.5 (0.1)
1.3 (0.3)
15 (3)
157a (87833)
0.095
0.005

0.2
0.014
0.017
0.07
0.3 (0.1)
2.5 (0.1)
20 (2)
56b (4671)
0.047
0.001

0.4
0.032
0.001
0.025
5.3 (0.8)
2.9 (0.2)
31 (3)
35c (2560)
0.013
0.007

0.3
0.020
0.005
0.05
0.8 (0.1)
2.7 (0.6)
15 (1)
34bc (2186)
0.030
0.002

lengths) (Webster & Ehrman, 1996). This concept can

be applied to cave streams and is particularly important in caves which are often nutrient polluted (e.g.,
Simon & Buikema, 1994; Boyer & Pasquarell, 1996).
Like inorganic nutrients, carbon spirals in streams,
but is also lost from the system as CO2 from respiration. Because of this loss, uptake length is not
a useful measure, and estimates of stream ecosystem efficiency, the extent to which energy inputs to a
stream are used within the stream (Webster & Meyer,
1997a), are more commonly used. Again, this concept
is applicable and useful in the study of cave streams.
Ecosystem efficiency is particularly interesting for
caves because it can be used to measure energy limitation and efficiency in what are considered energy-poor
ecosystems.
In this study, we examined ammonium retention
and whole stream metabolism in cave streams to: (1)
test the utility of standard surface stream methods and
models in cave streams, (2) estimate basic ecosystem
parameters for cave streams to provide a benchmark
for comparison with surface and cave streams, and (3)
begin to understand nutrient spiraling and carbon flow
in cave streams.

Methods
Study sites
The study streams were located in the Organ Cave
drainage basin, Greenbrier County, West Virginia. Organ Cave is large (>60 km) and contains a network of
streams ranging in size from small tributaries draining

the interface zone (epikarst) between soils and bedrock to large streams in the deep subsurface (Stevens,
1988). Organ cave streams contain a rich aquatic fauna
that has been the subject of previous studies examining evolutionary, population and community ecology
of the system (Culver et al., 1994). We measured nutrient uptake in 4 streams, Jones Canyon (JC), Lipps
Entrance (LE), Skid Row Side (SRS) and Sively 3
(S3), in March 1999 and whole-stream metabolism in
S3 in January 2000. All three streams are small (Table
1), contain little coarse organic matter and drain the
epikarst.
Nutrient uptake
We used short-term ammonium and conservative
tracer releases to measure nitrogen uptake and hydrologic properties of each stream (Webster & Ehrman,
1996). We chose ammonium because it was the least
abundant (Table 1) and most likely limiting nutrient. A solution of chloride, as a conservative tracer,
and ammonium chloride were injected to the stream
using a fluid metering pump at the top of a 30 to
75-m reach in each stream. The pump continuously
dripped solution until the ammonium concentration
in the stream plateaued at roughly 24 times background concentration at the release point. During and
after the release, we monitored stream chloride concentration using conductivity meters at the end of the
study reach to quantify discharge and velocity (Stream
Solute Workshop, 1990). The conductivity meters
were calibrated against standard chloride solutions to
translate conductivity to chloride concentration. When
chloride concentration reached plateau, triplicate wa-

33

Figure 1. Plot of regressions of ln transformed NH4 + :Cl ratios versus distance downstream from the injection point used to calculate NH4 +
uptake parameters. JC = Jones Canyon stream, LE = Lipps Entrance stream, SRS = Skid Row Side stream, and S3 = Sively 3 stream.

ter samples were collected from 4 to 8 points along the

stream reach and immediately passed through glass
micro-fiber filters (Whatman GF/F). Duplicate water
samples were collected at each station prior to the release to measure background nutrient concentration.
Plateau and background water samples were returned
to the laboratory on ice and analyzed for NH4 + N, and
NO3 N concentration using a Technicon Autoanalyzer. Soluble reactive phosphorus (SRP) in each
sample was measured using the ascorbic acid method
and a spectrophotometer equipped with a 10-cm cell
(APHA, 1995).
Ammonium uptake length was calculated based
on downstream changes in ammonium concentration
after correction for dilution using downstream changes
in chloride concentration. Uptake length, the average
distance traveled by an NH4 + N atom before being
removed from the water column, was calculated as:
Sw =

1
,
kc

where Sw is the uptake length (m) and kc is the

uptake rate coefficient (m1 , the slope of the ln
transformed regression of NH4 + N:Cl ratio versus
distance downstream, Fig. 1).
The ammonium mass transfer coefficient (vf ,
m/s), a measure of ammonium uptake efficiency that
normalizes for differences in discharge among streams

(Davis & Minshall, 1999), was calculated as:

vf =

uh
,
Sw

where u is the water velocity (m/s), h is the stream

depth (m), and Sw is the ammonium uptake length
(Stream Solute Workshop, 1990). Ammonium uptake rate, the amount of ammonium removed per unit
streambed area and time, was calculated as:
U = vf C,
where U is the uptake rate (g NH4 + N m2 min1 ),
vf is the mass transfer coefficient (m/min), and C is the
background NH4 + N concentration (g/m3) (Stream
Solute Workshop, 1990).
Metabolism
We measured whole-stream metabolism in one stream,
Sively 3, using the upstream-downstream dissolved
oxygen technique (Marzolf et al., 1994) with modifications suggested by Young & Huryn (1998). The
method involves calculating whole-stream metabolism
by measuring change in dissolved oxygen (DO) in a
packet of stream water as it moves along a stream
reach. Changes in oxygen content of the water depend
on primary production, respiration and gas exchange
with the atmosphere. Because there is no primary production in cave streams, only respiration and oxygen
diffusion, the reaeration flux, at the airwater interface

34
were relevant. Reaeration flux depends on the reaeration coefficient of oxygen (koxygen) and the oxygen
deficit of the water (DOdeficit ). We used propane as
a surrogate gas to measure koxygen and chloride as a
conservative tracer to calculate discharge, water travel
time between sites ( ) and propane dilution caused by
water entering the stream along the study reach.
We injected chloride, using a fluid metering pump,
and propane, using a propane cylinder and diffuser, at
the top of a 100-m reach of Sively 3. Chloride concentration was monitored at the bottom of the reach and
once chloride reached a steady state, water samples
were collected at the top of the reach. Water was then
collected at the bottom of the reach one travel-time
( ) after the upstream samples, so the same packet of
water was sampled at the upstream and downstream
stations. Travel time was calculated as the time to
one half chloride plateau concentration (Marzolf et al.,
1994). At each station, triplicate water samples were
collected in syringes and immediately injected into vacuum tubes, leaving a headspace at the top of the tube.
The vacuum tubes were returned to the laboratory on
ice and propane in the headspace was measured using
a gas chromatograph. At the time of water collection,
dissolved oxygen (DO) was measured at both stations
using a YSI dissolved oxygen meter.
The propane exchange coefficient, kpropane (min1)
was calculated as:



G1 C2
k propane = 1 ln
,
G2C1
where is the reach travel time (min), G1 and G2
are the propane concentrations at the upstream and
downstream stations, respectively, and C1 and C2
are the steady state chloride concentrations at the upstream and downstream stations, respectively. Using
this, koxygen was calculated as 1.39 kpropane (Marzolf et al., 1994). Reaeration flux (RF, g/L) was then
calculated as:
RF = DOdeficit koxygen ,
where DOdeficit is the difference between 100% saturation and the measured DO concentration in the stream
(g/L), koxygen is the oxygen exchange coefficient
(min1), and is the reach travel time (min).
Community respiration (CR, g O2 m2 s1 ) was
calculated as
(DORF)Q
,
CR =
A
where DO is the difference between upstream and
downstream DO concentration (g/L), RF is the reaeration flux (g/L), Q is discharge (L/s), and A is the

reach area (m2 ). We converted CR to units C respired

using a respiration quotient of 0.85 (Bott, 1996). We
also calculated specific respiration (SR), i.e., respiration rate per g benthic organic carbon (BOC), and
BOC turnover time (BOC standing stock/CR).
Benthic organic matter
We used measures of coarse and fine particulate organic matter (CPOM and FPOM, respectively) standing stocks from collections on 24 March 1998. These
samples were collected at 5-m intervals along a 50-m
reach of each stream. CPOM was collected by placing a Surber sampler (0.09 m2 , 1-mm mesh) on the
stream bottom and stirring the substrate by hand to
wash CPOM into a net. FPOM was collected by pushing a 10-cm diameter cylinder 5 cm into the substrate
and sealing around the cylinder with foam insulation.
Sediment and water in the cylinder were stirred, the
slurry was removed using a hand pump, and triplicate subsamples were collected and frozen. CPOM was
air dried, sorted in the laboratory into wood and other
particles, then ground in a Wiley mill. Subsamples of
the FBOM slurry were filtered onto pre-ashed Gelman
AE filters. FPOM and ground CPOM samples were
dried at 60 C, weighed, ashed at 550 C, wetted with
deionized water, dried at 60 C and re-weighed to
calculate ash-free dry mass (AFDM).
Statistical analyses
Uptake lengths were compared among streams using
analysis of covariance (ANCOVA) with a dummy variable technique (Kleinbaum et al., 1988). Correlations
among uptake parameters (Sw , vf and U ), nutrients (NH4 and NO3 ), discharge and benthic organic
matter standing stock were determined using Pearson Product Moment Correlation coefficients (Sokal
& Rohlf, 1995). Literature values for ammonium uptake lengths and discharge for surface streams were
collected and analyzed using linear regression for
comparison to cave streams.

Results
Nutrient uptake
Nitrate and SRP concentrations were high in the cave
streams while ammonium was quite low (Table 1).
Ammonium was high in LE as compared to the other

35
Table 2. Benthic organic matter standing stocks, community and
specific respiration, and benthic organic carbon turnover time
for cave and surface streams. Surface stream values are means
with ranges in parentheses. CPOM = coarse particulate organic
matter. FBOM = fine benthic organic matter. CR = community
respiration. BOC = benthic organic carbon. = not measured
Parameter

S3

JC

LE SRS

Surface Streams

CPOM (g AFDM/m2 )

0.09 0 41.4 0 6121 (2.734 090)a

FPOM (g AFDM/m2 )
28.2 37.4 48.6 44.3
346 (01903)a
Total BOM (g AFDM/m2 ) 28.3 37.4 90.0 44.3 7667 (2035 157)a
CR (g C m2 y1 )
32.9

265 (10 1200)b

Specific respiration (y1 ) 2.6

1.2 (0.00211.848)b
BOC turnover (y)
0.4

55.1 (0.1 425.0)b

a Data from Webster & Meyer (1997).
b Data from Sinsabaugh (1997).

streams (Table 1). S3 was larger and had higher discharge and velocity than JC, LE and SRS (Table 1).
Ammonium concentration declined downstream during the releases, yielding uptake lengths ranging from
34 to 157 m (Fig. 1, Table 1). Uptake length was
significantly higher in S3 than in the other streams
which had similar uptake lengths (Table 1). Ammonium uptake length was positively correlated with
discharge (p = 0.006, r = 0.99) and negatively correlated with nitrate concentration (p = 0.016, r =
0.98). Normalization of uptake length by velocity
and depth only slightly reduced differences in uptake among streams; mass transfer coefficients (vf )
ranged from 0.013104 to 0.095104 m/s and
were more than two times higher in S3 than in the
other streams (Table 1). Ammonium mass transfer
was negatively correlated with nitrate concentration
(p = 0.016, r = 0.98). Uptake rate (U ) ranged
from 0.001 to 0.007 g N m2 s1 and was not correlated with any measured parameters. Discharge was
negatively correlated with nitrate concentration (p =
0.020, r 2 = 0.98). Benthic organic matter standing
stock was not significantly correlated with any other
parameters.
Benthic organic carbon and metabolism
Most of the organic matter in the cave streams was
FPOM (Table 2). FPOM standing stocks in the cave
streams fell in the low range of values reported for
surface streams (Table 2). Amount of CPOM in 3 of
the cave streams (JC, SRS and S3) was well below
amounts reported for surface streams (Table 2). Standing stock of CPOM in LE was within the low range of
CPOM reported in surface streams.

Discharge and temperature during the metabolism experiment were 0.4 L/s and 6.7 C, respectively.
Sively 3 is a well-oxygenated, turbulent stream (>95%
saturation). Changes in downstream propane concentration resulted in a koxygen of 48.4 d1 . Community
respiration was 32.9 g C m2 y1 , which, when combined with BOC standing stock, resulted in a specific
respiration of 2.6 y1 and a BOC turnover time of 0.4
y.

Discussion
Nutrient uptake
Nutrient uptake length depends on stream velocity and
depth as well as biological nutrient demand (Newbold, 1992). High discharge, and correspondingly
high velocity and depth, increases the rate of downstream nutrient transport, thereby increasing nutrient
uptake lengths (DAngelo & Webster, 1991; Valett et
al., 1996). In our survey of published values for ammonium uptake lengths in surface streams, there was
a strong positive relationship between discharge and
uptake length (Fig. 2). This relationship between ammonium uptake length and discharge has been noted
by other authors (Butturini & Sabater, 1998; Peterson
et al., 2001). The positive relationship between uptake
length and discharge in the cave streams suggests that
discharge is an important factor in determining ammonium transport in cave streams as is the case in
surface streams.
Considering the absence of primary producers and
low organic matter standing stocks in caves, we expected long nutrient uptake lengths and low nutrient
demand in cave streams. Because discharge in JC,
LE and SRS was lower than in any surface streams
with reported ammonium uptake lengths we cannot
directly compare our streams to surface streams of
similar size. However, the cave streams were much
less ammonium retentive, i.e. they had longer uptake lengths, than would be predicted considering their
small size and the relationship between uptake length
and discharge (Fig. 2). Uptake length in S3 was longer
than all other streams of equivalent discharge (Fig. 2).
Kopcek & Blazka (1994) reported discharge (1.12.2
L/s), ammonium uptake lengths (82190 m) and vf
(0.11040.19104 m/s) in a surface stream that
were similar to our measurements in S3 (Fig. 2). Like
our cave streams, Kopcek & Blazkas (1994) stream
had relatively long uptake lengths for its size (Fig. 2).

36

Figure 2. Ammonium uptake length versus stream discharge for cave streams (JC, LE, SRS and S3) and surface streams (110). Solid line is
the regression of uptake length versus discharge for surface streams (log Sw = 0.567(log Q) + 1.089, p < 0.0001, r 2 = 0.74). Dashed lines are
95% prediction intervals for the regression. JC = Jones Canyon stream, LE = Lipps Entrance stream, SRS = Skid Row Side stream, and S3 =
Sively 3 stream. 1 = Kopacek & Blazka, 1994, 2 = Webster et al., 2000, 3 = Wallace et al., 1995, 4 = Tank et al., 2000, 5 = Hall et al., 1998, 6
= Marti & Sabater, 1996, 7 = Peterson et al., 2001, 8 = Wolheim et al., 2001, 9 = Mulholland et al., 2000, and 10 = Dodds et al., 2000, 11 =
Butturini & Sabater, 1998.

Their stream was at high elevation, acidic (pH 4.50

4.65) and surrounded by sparse alpine meadows that
likely contributed little coarse organic matter to the
stream. These factors may have reduced biological
ammonium demand, resulting in a surface stream that
was functionally more similar to our cave streams than
to other surface streams with higher nutrient demand.
Because Sw is strongly influenced by discharge,
mass transfer coefficients, which account for differences in velocity and depth, are a better measure of
biological nutrient demand for comparisons among
streams (Davis & Minshall, 1999). Ammonium mass
transfer coefficients were low in the cave streams
(0.0301040.095104 m/s) as compared to surface streams (range = 0.11047.0104 m/s,
Kopcek & Blazka, 1994; Sabater et al., 2000;
Peterson et al., 2001), indicating low biological ammonium demand in the cave (Table 1). Low ammonium demand in the cave was also reflected in the
low ammonium uptake rates in the cave (0.0010.007
g N m2 s1 ) compared to surface streams (0.163.8
g N m2 s1 , Kopcek & Blazka, 1994; Sabater et
al., 2000; Webster et al., 2000; Peterson et al., 2001).
Overall low nutrient retention in the cave streams as
compared to surface streams is probably a result of low
amount of organic matter in the cave streams. Litter
exclusion and wood removal reduce N and P retention

in surface streams (Webster et al., 2000). Cave streams

are extreme examples of streams disconnected from
detrital input by riparian vegetation.
While differences in uptake length among cave
streams can be reasonably explained by differences
in discharge among the streams, differences in mass
transfer coefficients, which normalize for discharge,
among cave streams suggest underlying differences
in biological demand among the streams. Because
primary producers are absent, assimilatory uptake by
heterotrophs and nitrification should be the only biotic
factors responsible for ammonium retention. Given the
low amount of benthic organic matter and the ample
supply of N and P in the cave streams, assimilatory
uptake by hetertrophic microbes may not play as great
a role in nutrient uptake in cave streams as in surface
streams (e.g., Tank et al., 2000). Indeed, ammonium
retention in the cave streams was not related to BOM
standing stocks and previous research showed little
evidence of N or P limitation of wood biofilm biomass
or respiration in these streams (Simon & Benfield,
2001). Ammonium demand by microbial films on detritus in cave streams is likely low relative to nutrient
availability.
Nitrification can account for a large portion, up to
60%, of ammonium uptake in some surface streams
(Peterson et al., 2001). Very low ammonium and high

37

Figure 3. Log10 BOC standing stock (A), log10 heterotrophic respiration (B), log10 specific respiration (C), and log10 BOC turnover time
(D) as functions of mean annual water temperature for surface streams (open symbols) and January and mean temperature for Sively 3 (filled
symbols). Regression lines are for surface streams only. Surface stream data from Sinsabaugh (1997).

nitrate concentrations suggest nitrifiers may be active

in the cave streams. Peterson et al. (2001) found that
surface streams with high nitrate levels usually had
high nitrification rates. Ammonium uptake length was
shorter in cave streams with high nitrate concentration,
suggesting nitrification rates may have influenced ammonium retention. However, ammonium mass transfer, a better indicator of biological retention, was lower
in cave streams with high nitrate concentration, which
is the opposite of what should occur if nitrification
was dictating ammonium retention. The relationship
between nitrate concentration and ammonium uptake
length is probably a result of positive correlation of
both factors with discharge. Any influence of nitrification rate on cave stream nitrate concentration is
probably small compared to surface sources (e.g. agriculture) and the large amount of nitrate often seen in
cave streams (Boyer and Pasquarell, 1996).
Ammonium also adsorbs to sediment (Richey et
al., 1985), especially fine particles, and differences
in adsorption could have caused differences in ammonium uptake among streams. Substrate particle size
distribution among the streams is similar (Simon &

Benfield, unpublished data) but we cannot confidently

discount differences in abiotic adsorption among the
cave streams.
Carbon processing
In a survey of 26 surface streams, Sinsabaugh (1997)
found BOC varied widely across biomes (Table 2, Fig.
3). Relative to those surface streams, BOC standing
stocks were quite low in the cave streams. The surface streams most similar to our streams were the
Kuparak River and Monument Creek (Alaska) that
have low detritus input (Harvey et al., 1997; Irons
& Oswood, 1997). Like these Alaskan streams, the
primary cause of low organic matter in our streams is
the lack of CPOM (Table 2). Low CPOM in our cave
streams is a result of the absence of vegetation along
the stream channel and constraints placed on detritus
input from the surface. JC and SRS are fed by percolating water and have no direct entrances large enough
to allow CPOM to enter the streams. The S3 stream
does have an opening to the surface but vegetation
has been cleared from the entrance and, consequently,

38
only a small amount of CPOM is washed underground
and into the stream. LE, which had a relatively large
amount of CPOM, had intact vegetation around a large
sinkhole entrance.
Compared to most surface streams, community
respiration was relatively low in S3 (Table 2, Fig. 1),
but S3 does fit well in the trend of respiration versus
mean annual stream temperature for surface streams
(Fig. 1). Surface streams with metabolic rates most
similar to S3 are in grasslands and boreal and montane coniferous forests. The grassland stream has low
standing stocks of BOC (Gray, 1997), and the coniferous forest streams are heavily shaded and have low
temperatures (Webster & Meyer, 1997b); S3 shares
both of these attributes.
Based on carbon turnover time, S3 is a relatively
carbon limited stream. Fast BOC turnover and high
specific respiration were the result of CR similar to
surface streams combined with low BOC standing
stock (Fig. 1). Low standing stock of wood, in particular, contributes to the rapid BOC turnover time in
S3. Turnover times are long in surface streams that
contain large amounts of wood, which decays slowly
in streams (Webster & Meyer, 1997a). Most organic
matter in the stream bed of S3 is FPOM which is used
more rapidly. In a previous study, leaves placed in
S3 broke down rapidly, also suggesting the stream is
carbon limited (Simon & Benfield, 2001).

Conclusions
Cave streams represent a heterotrophic extreme in the
continuum of stream types. The concepts and methods
of nutrient and carbon spiraling developed for surface streams are applicable and useful in the study of
cave streams. Ammonium transport in cave streams
is predominantly determined by discharge, but biological factors may be involved in ammonium retention.
Given the low organic matter supplies and aerobic
conditions, nitrification may be an important factor in
N cycling in cave streams. The roles of nitrification
and abiotic adsorption need to be explored in cave
streams. Ammonium demand in the cave streams was
very low and the stream community does not seem
to be nitrogen limited. As a result, excess nitrogen
from human sources is not likely to be retained in cave
drainages. Cave streams therefore have little capacity
for remediating nitrogen pollution in karst areas.
Unlike nitrogen, carbon appears to be limiting in
at least some cave streams. BOC is cycling rapidly

through S3, suggesting organic matter is in high demand. This stream has been disconnected from surface
vegetation by human activity, but contains an aquatic
community more typical of streams in the drainage
that receive detritus input. Cave streams with and
without natural input need to be studied and compared to further understand energy limitation and the
influence of land use on cave streams.

Acknowledgements
We thank S. and J. Morgan for allowing us access
to Organ Cave through their property. S. Thomas assisted with the metabolism field measurements. This
research was supported by a National Science Foundation Dissertation Improvement Grant, DEB-9801082
and supporting grants from the National Speleological
Society, Sigma Xi, and the Biology Department and
Graduate Student Association at Virginia Tech.

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