Ecological Modelling 137 (2001) 61 – 75

www.elsevier.com/locate/ecolmodel

A model of the development of a periphyton community:

resource and flow dynamics
Takashi Asaeda *, Duong Hong Son
Department of En6ironmental Science and Human Technology, Saitama Uni6ersity, 255, Shimo-okubo, Urawa,
Saitama 338 -8750, Japan

Received 16 February 2000; received in revised form 11 September 2000; accepted 6 October 2000

Abstract

An ecological model of periphyton dynamics was designed to approach the development of a periphyton
community under the effects of surrounding nutrient concentration and light intensity. The model simulated the
vertical heterogeneities of biomass, light irradiance, and nutrients within the periphyton mat. To determine the
competition for resources between species, a Monod-type relationship was used to relate external nutrient concentra-
tion to periphyton uptake rate, and ‘Liebig’s law of the minimum’ was used to combine light and nutrient effects on
growth rate and tension strength of the algal cell. The effect of the periphyton community to the vertical distribution
of water velocity was also numerically investigated. To confirm the validities, the developed model was verified using
several cases: the external concentration was constant or was varied with different diffusion coefficients. Reasonable
responses of the periphyton community to the ambience nutrient concentrations, light intensity and, in turn, temporal
variation of nutrients during the development of a periphyton community strongly encouraged further application of
the model. Variation in the water velocity in a vertical direction intensively contributed to the results of the model.
© 2001 Elsevier Science B.V. All rights reserved.

Keywords: Algal competition; Nutrient uptake; Periphyton; Photosynthesis

1. Introduction dynamics of the essential resources for periphyton

Studies of stream ecosystems have focused on
periphyton communities, because benthic algae
usually dominate planktonic communities. Many
experimental studies have concentrated on the
* Corresponding author. Tel.: +81-48-8583563; fax: + 81-
48-8583563.
E-mail address: asaeda@env.civil.saitama-u.ac.jp (T. As-
aeda).
growth. Although several studies have been made
on the photosynthesis –irradiance relationship
(Sand-Jensen and Revsbech, 1987; Boston and
Hill, 1991; Dodds, 1991; Graham et al., 1995;
Hill, 1996), only a few studies have evaluated the
relation between light inside a matrix and the
algal response.
The relation between periphyton and nutrients
has been intensively studied. Borchardt (1994)
found different optimum N:P ratios for Spirogyra

0304-3800/01/$ - see front matter © 2001 Elsevier Science B.V. All rights reserved.
PII: S0304-3800(00)00432-4
62 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75
flu6iatilis at different velocities. Newbold et al.
(1981, 1983) examined the horizontal heterogenei-
ties of nutrients due to a spiraling process using
both models and experiments. In natural streams,
the saturation concentration of nutrients on a
periphyton community seems to depend strongly
on the density of the community (Bothwell, 1985,
1988, 1989).
Several models have been devised to clarify the
relations among the periphyton community, nutri-
ent concentration in the overflowing water, and
the nutrient availability for periphyton growth
(Kim et al., 1992; Mulholland et al., 1994; DeAn-
gelis et al., 1995). The vertical variation of oxygen
produced in photosynthesis (Carlton and Wetzel,
1987; Sand-Jensen and Revsbech, 1987; Carlton
and Wetzel, 1988; Bott et al., 1997) implies a
vertically varying concentration of nutrient re-
sources for periphyton growth, indicating the pos-
sibility of a nutrient-limited condition for
periphyton communities (Horner et al., 1990;
Dodds, 1991; Borchardt, 1996). However, the ver-
tical variation of the resources for periphyton
growth, such as light and nutrients, has not been
quantitatively examined by either models or
experiments.
As a number of problems have been pointed
out in experiments, such as altering the commu-
nity structure and producing higher uptake rates
by removing the community from the surface in
the experiment (Kim et al., 1992), in this study we
used models to: (1) simulate the process of periph-
yton development under the effects of essential
resources such as nutrients and light; and (2) in
turn, model the resources and water flow in the
development of the periphyton community.
2. Model description
All processes are given in the appendix to com-
pute the thickness of the periphyton mat, light
resource, growth, settlement and detachment of
periphyton (Asaeda and Son, 2000). Table 1 lists
the symbols.
An important concept underpinning algal re-
source kinetics is the single resource limitation
(Hamilton and Schladow, 1997), or the growth of
a species being limited by only one resource at a
time, ‘Liebig’s law of the minimum’. Thus, the
resource index, which is defined as the indicator
of the health condition and the reproduction abil-
ity of the cell, is given as:
Rn, k, i( j ) = min[NRn, k, i( j ), IRn, k ,i( j )] (1)
Although the multiplication form of these compo-
nents is an alternative for the resource index, Eq.
(1) seems to provide better results (Haney and
Jackson, 1996). Particularly, the multiplication
form becomes problematic when a number of
resources increase.
Some periphytic algae grow within the viscous
sub-layer, where the flow is substantially laminar
(Horner et al., 1990); however, filamentous algae
are found mainly outside the viscous sub-layer
(Dodds and Gudder, 1992). Fig. 1 shows the main
processes in the water column and their directions
of influence.
2.1. Detritus
The amount of detritus is simply assumed to be
produced only from dead cells in a periphyton
mat, and is given as a result of mortality, decom-
position, and detachment as:
(DCk, i( j)
= % GmortalCn, k, i( j)
(t n
− (q T − 20Gdecomp + Gdetach)DCk, i( j)
(2)
2.2. Nutrients
Although the process of the internal nutrient
concentration during reproduction is biologically
greatly complex, the changing rate is thought to
depend on the reproduction rate (Haney and
Jackson, 1996). In the reproduction process of a
cell, some nutrients inside the parent cell are
assumed to conservatively succeed to new daugh-
ter cells. Thus
(IN · C)n, k, i( j) = (INnew · Cnew)n, k, i( j) (3)
If the reproduction rate is a function of tempera-
ture, maximum reproduction rate, and resource
sufficiency (Eq. (4A)), excluding the subscripts
from Eq. (3) yields:
 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75 63

Table 1 Table 1 (Continued)

Glossary of symbols
Symbol Unit Definition
Symbol Unit Definition
UNmax mg cell−1 day−1 Maximum rate of nutrient
B mm3 mm−2 Total biomass uptake
C cell mm−2 Cell density Vi m3 Biovolume of filamentous cell
Cd Drag coefficient Z m Distance from the bottom
DC cell mm−2 Dead cell density Dz m Thickness of k-layer
EN mg l−1 External nutrient pw, pc m−1 Abiotic and biotic attenuation
concentration constant
ENo mg l−1 Upstream nutrient z, za kg m−3 Water, algal specific density
concentration (boundary ~bottom N m−2 Bottom shear stress
condition) ( =zfU 2/H)
f Bottom friction coefficient
Fcr N Threshold strength of the
filament IN · C=INnew · Cnew = INnew(C+DC)
g m s−2 Acceleration due to gravity 6
Gdetach day−1 Detachment rate for =INnew(C+Cq T − 20GdivRDt) (4)
non-filamentous species
−1
Gdiv day Reproduction rate or:
Ggz day−1 Grazing rate
Gmortal day−1 Mortality rate IN
Gres day−1 Respiration rate
INnew = T − 20
(5)
1+ q GdivRDt
H m Depth
Ik mE m−2 s−1 Light irradiance at k-layer Thus, the loss rate of the internal nutrient concen-
IN mg cell−1 Internal nutrient tration due to the reproduction is given by
concentration of the cell
INmin mg cell−1 Minimum internal nutrient (IN INnew − IN INq T − 20GdivR
= =− (6)
concentration for (t Dt 1+ q T − 20GdivRDt
reproduction
Isat mE m−2 s−1 Saturation irradiance for In this study, the reduction of internal nutrient
reproduction concentration due to the respiration process is
K Von Karman constant assumed simply as a function of the respiration
(= 0.4)
Kdiff m2 s−1 Diffusion coefficient
rate, the internal nutrient concentration, and the
KN mg l−1 Half-saturation nutrient temperature as GresINq T-20 (Hamilton and
concentration for the uptake Schladow, 1997).
process Benthic algal communities usually comprise an
Kvis m2 s−1 Viscosity coefficient outer layer of actively growing cells that remove
Kz m s−2 Calibration constant
l m Longitudinal distance
nutrients from the overflowing water and an inner
L cell filament−1 Length of filament layer of older, metabolically inactive cells (Mul-
n,k,i( j ) Day, layer, and filamentous holland, 1996). In the study, however, all cells of
(non-filamentous) species a species in each layer were assumed to have the
index same characteristics in nutrient uptake. The inter-
Ndetach filament day−1 Detachment rate for
nal nutrient concentrations and the maximum up-
filamentous species
NR, IR Nutrient and light resource take rates were selected according to the size of
index algal species (Borchardt et al., 1994).
Q Temperature constant The nutrient uptake rate of a cell is assumed to
SI m2 Surface area of filamentous correlate inversely with the internal nutrient con-
cell
centration, being zero when the cell saturates, and
Snor m−2 Trapping efficiency
T °C Water temperature to positively correlate with the nutrient con-
t day Time centration in the surrounding water by a Michae-
U m s−1 Water velocity lis-Menten-type relationship (Hamilton and
64 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75

Fig. 1. A schematic diagram shows the main processes inside the periphytic mat. Filled boxes are simulated constituents, rounded
boxes are related processes, hexagon and pentagons are controlling factors, and arrows are influencing directions.

Schladow, 1997). Endogenous requirements for Overflowing water exchanges nutrients continu-
nutrients are the main cause of thresholds for ously with the periphyton mat. The diffusion pro-
growth, and thus the minimum concentration of cess through the top boundary layer of the
internal nutrients is often defined as the concen- periphyton mat, regardless of molecular or turbu-
tration to stop the reproduction. Combining these lent diffusion, controls the supply of nutrients
relations with the losses due to reproduction and markedly (Mulholland et al., 1994). If nutrient
respiration provides the expression for the ex- exchanges from the sediment and other processes
change rate of the internal nutrient concentration: were negligible, the conservation of the external
nutrient concentration in the k-layer is given as:
(INn, k, i( j)
(t (ENk (ENk ( 2ENk
+U =Kdiff
IN
= UNmaxi( j)q T − 20 maxi( j)
−INn, k, i( j) ENk (t (x (z 2
INmaxi( j) −INmini( j) KNi( j) +ENk
10 − 3 T − 20
IN q T − 20Gdivi( j)Rn, k, i( j) + q Gdecomp % DCn, k, i( j)INn, k, i( j)
− n, k, i(Tj)− 20 DV
1+q Gdivi( j)Rn, k, i( j)Dt i( j)

− GresINn, k, i( j)q T − 20 (7) 10 − 3

− % % UNmaxi( j)q T − 20
The nutrient resource index is defined as DV n i( j)

INn, k, i( j) −INmini( j) INmaxi( j) − INn, k, i( j) ENk

NRn, k, i( j) = (8) C (9)
INn, k, i( j) INmaxi( j) − INmini( j) KNi( j) + ENk n, k, i( j)
 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75
The ratio of 10 − 3/DV is used to convert the
uptaken and decomposed nutrient rate (mg day − 1)
to the exchange rate of the nutrient concentration
(mg l − 1 day − 1) in the layer. In Eq. (9), the left
terms indicate the unsteadiness and the advective
transport of the nutrient concentration, and the
right terms are the diffusion, the decomposition of
detritus, and the loss due to algal uptake. The left
hand side and the first term on the right hand side
are well-defined for mass transport in fluid me-
chanics and the remaining terms are sources and
sinks inside the layer.
Although excluding nutrients released from the
substrate, the model considers nutrient cycling
through dead cells (detritus). This assumption is
satisfactory for most laboratory experiments on
periphyton development, where the substrate is
usually made from ceramic, plastic, and clay tiles
and no nutrients are released from bottom. Fur-
thermore, nutrient exchanges in nature between
the sediment and over-flowing water are so com-
plicated that they may be represented by another
independent sub-model (Asaeda and Bon, 1997).
If nutrient flux from the air and from the
sediment is assumed to be negligible, the
) )
boundary conditions for Eq. (9) are:
(EN (EN
=0 =0 (10)
(z surface (z bottom
Provided that the amount of a nutrient is con-
served in the water column regardless of its forms
and that the main processes in the nutrient ex-
change occur as in Fig. 1, Eqs. (1), (7) – (10) are
repeatedly applied for all concerned nutrients. In
this study, these equations were used to evaluate
temporal variations of phosphorous and nitrogen
concentrations. More detailed processes in the
cycling of the specific nutrient can be coupled to
the governing Eqs. (7) and (9).
2.3. Flow dynamics
The algal mat lies from the viscous sublayer
through the overlying outer boundary layer, and
thus, the local Reynolds number varies widely. If
a steady and longitudinally homogeneous current
is assumed, the friction at the bottom and the
drag force on the periphyton mat balances with
the longitudinal pressure gradient as:
65
1 (P ( 2U B 2/3
0= − + Kvis 2 − CdU 2 (11)
z (x (z DV
where (P/z(x is the longitudinal pressure gradi-
ent and is given by the vertical integration of Eq.
(11):
1 (P ~bottom B 2/3
= − C dU 2 (12)
z (x z DV
The last term in Eq. (11) represents the resistance
due to the periphyton mat, and is similar to the
equation used for a terrestrial plant canopy
(Kondo and Watanabe, 1992).
Inside the viscous sublayer, the kinetic viscosity
of water and the linear velocity distribution were
used. Outside the viscous layer, turbulent viscosity
and diffusivity were calculated from the velocity
) )  
profile and the mixing length, lm, as:
(U z 1/2
Kvis = Kdif = l 2m where lm = Kz 1−
(z H
(13)
3. Numerical procedure
A system of differential equations was solved
numerically by coding in FORTRAN90, includ-
ing 23 subroutines of  2300 command lines.
Additional graphic subroutines authorized by Mi-
crosoft corporation permitted the main program
to simultaneously show the results. The model
was also programmed with highly flexible running
options. For accuracy, the fourth-order Runge-
Kutta method was initially used to solve a system
of the first-order differential equations. To in-
crease the computational stability of the diffusion
process and to save computation time, an implicit
scheme was finally used for the external nutrient
concentrations and vertical profile of velocity. A
Thomas algorithm was used to solve a tri-diago-
nal matrix (Fletcher, 1991), while an explicit
scheme was used to integrate all the remaining
variables. The stability condition of the vertical
diffusion, Dt BDz 2/4Kdiff, was satisfied (Vreug-
denhil, 1994). Thus, a few seconds of the time step
was used in our model, while several hours of
time step was used in previous models (EPA,
1985). To deal with high biomass environments a
66 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75

small time step is a prerequisite condition to reduce

where EN0 is the boundary condition for nutrients
errors caused by numerical integration. If a large
and l is the length of a stream. In this aspect,
time step was chosen, for instance, a dense periph-
considering one more dimension (e.g. longitudinal)
yton in a thin layer would exhaust the whole
certainly provides results closer to reality than this
nutrient even within a period less than the time step.
study.
The velocity was calculated upwards immedi-
ately from the viscous sublayer, while the interpo-
lation of linear velocity distribution was adopted
4. Verification and discussion
inside the boundary layer. The water depth was
divided into a number of layers, their thickness
In the verification, the major nutrients of concern
gradually increasing from the bottom to the water
were nitrogen, and phosphorous because they are
surface. A resolution of one-third of the sublayer
essential for aquatic organism. Other nutrients,
thickness was always applied to the bottom layer.
such as carbon and silicon, were assumed to be
The vertical resolution of the overlying mesh was
sufficient for periphyton growth. Boundary values
strongly related to the hydraulic conditions. In each
of the phosphorous and nitrogen concentrations
layer, the relationships of the periphyton biomass,
(e.g. EN0 in Eq. (9)) were tentatively assigned at 2
internal and external nutrient concentrations, irra-
and 15 mg l − 1, respectively. The water depth,
diation, and the hydraulic regime of the overflow-
depth-average water velocity, and the light intensity
ing water were established.
at the water surface were assumed to be constant
Although this study is one dimension (vertical),
as 6 cm, 20 cm s − 1, and 150 mE m − 2 s − 1,
the advective transport of nutrients can not be
respectively. Table 2 lists the biological parameters
neglected because nutrients in a water column are
of filamentous and non-filamentous species. The
limited compared with the requirements of the
resource index for periphyton species was calcu-
periphyton community. If the position of a consid-
lated from Eqs. (1), (8) and (3A).
ered area is in the middle of a stream and the
nutrient gradient is linear along the stream, the
4.1. Constant external nutrient concentration
contribution of nutrients from a convection, the
second term of Eq. (9), is numerically represented
The first verification was made under constant
as
external nutrient concentrations or, in other words,
EN0 −ENk an infinite diffusion coefficient. The verification
U ,
0.5l consisted of three cases: (a) the conditions

Table 2
Biological parameters used in the model (nutrient-related parameters are for either phosphorus or nitrogen)

Symbol Unit Filamentous i-species Non-filamentous j-species

Gdecomp day−1 0.02 0.02

Ggz day−1 0 0
Gres day−1 0.1 0.1
INmin mgP cell−1 10×10−7 0.6×10−7
mgN cell−1 200×10−7 4.2×10−7
INmax mgP cell−1 70×10−7 4.2×10−7
mgN cell−1 1400×10−7 29.4×10−7
Isat mE m−2 s−1 200 200
KN mgP l−1 10.0 2.0
mgN l−1 30.0 4.0
UNmax mgP cell−1 day−1 1×10−5 0.05×10−5
mgN cell−1 day−1 14×10−5 0.35×10−5
 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75 67

Fig. 2. Cell density of filamentous (solid lines) and non-filamentous (dashed lines) species in three cases: a, b, and c (refer to the text).

Fig. 3. Temporal variation of the internal phosphorous concentration of filamentous (solid lines) and non-filamentous (dashed lines)
cell in the bottom layer in three cases: a, b, and c (refer to the text).

were as described in Table 2; (b) the minimum
and the maximum internal nutrient concentra-
tions of case (a) were reduced by 50%; and (c) the
nutrient uptake rates of all species of case (a) were
reduced by 50%. The time step of 60 s was used.
Fig. 2 shows the cell density of filamentous and
non-filamentous species in three cases. In case (a),
the peak biomass of filamentous and non-filamen-
tous species was 8000 and 20 000 cell mm − 2 on
days 19 and 17, respectively (Fig. 2, case (a)). At
the peak biomass, however, the non-filamentous
community was dislodged and its density gradu-
ally decreased, while the filamentous community
notably fluctuated because remaining filaments
detached intermittently. As these detached
filaments were considerably long, consisting of
many cells, their detachment markedly affected
the total density of the community.
The decrease in the maximum and minimum
internal nutrient concentrations by 50% slightly
shifted the peak values of both filamentous and
non-filamentous biomass to the earlier days (16
and 14), because the decrease in the maximum
and minimum nutrient concentration conse-
quently increases the uptake rate in Eq. (7) and
eventually the growth rate (Fig. 2, case (b)). Thus,
the periphyton community becomes more vulner-
able to the drags and resource depletion.
With decreasing uptake rates, the biomass of
non-filamentous species in case (c) did not in-
68 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75
crease as fast as cases (a – b). Consequently,
filamentous and non-filamentous biomass started
being dislodged later: on day 27 and day 16 (Fig.
2, case (c)). Especially, the extended high biomass
period of non-filamentous species delayed the in-
tensive growth of filamentous species by nearly 10
days.
Fig. 3 shows the variation of the internal phos-
phorous concentration, as an example of the in-
ternal nutrients concentration, where both
phosphorous and nitrogen had similar trends. The
value of a cohort settled at the beginning is repre-
Fig. 4. Temporal variation of the phosphorous (solid lines),
nitrogen (dashed lines), and light (dotted lines) resource index
of filamentous (left) and non-filamentous (right) cell in three
cases: a, b, and c (refer to the text). Arrows indicate detach-
ment of filaments.
sented for a non-filamentous species, whereas for
a filamentous species, the value of an individual
filament is used. Reducing concentration to zero
shows the detachment of concerned filament.
Notably, the internal nutrient concentration
was reduced markedly for both filamentous and
non-filamentous species within the first few days
after settling, because of the lower ambient nutri-
ent condition than the cell experienced before. As
the uptake rate depends on the ambient nutrient
concentration, if a cell settles on a nutrient-poor
substratum, the internal nutrient concentration
decreases rapidly. After that, a markedly stable
level of the internal nutrient concentration lasts
for both species (Fig. 3). As the external nutrient
remains constant regardless of the biomass, the
uptake rate of an individual cell balances with the
loss due to reproduction and respiration once the
internal nutrient concentration level satisfies this
condition until the limiting factor changes to the
irradiance. The markedly stable level of the inter-
nal nutrient concentration lasts for  15, 10, and
22 days in cases (a–c), respectively (Fig. 3). The
internal nutrient concentration increased slightly
before the filament detachment.
Fig. 4 shows the temporal variation of resource
indices of phosphorous nitrogen and light, in
these cases. Generally, when a species switches
from being nutrient-limited to light-limited, the
nutrient resource indices increase. In due course,
an increasing biomass consequently decreases the
light index (Eq. (3A)), growth rate (Eq. (4A)), and
the reproduction loss (Eq. (5)), and finally in-
creases the internal nutrient concentration (Eq.
(6)). Thus, immediately after the light condition is
limiting and further worsens with increasing
filamentous algal biomass, the internal nutrient
concentration recovers.
Fig. 4 also shows that filaments detach earliest
in case (b) and latest in case (c). A higher resource
index of a single cell implies a healthier filament
and, thus, the filament is more difficult to be
detached. The result indicates, however, that
filaments with a higher resource index were de-
tached sooner. The drag force increases exponen-
tially with an increase in resource index, while the
tension strength of a cell increases only linearly
with an increase in resource index. The biomass
 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75 69

required in the computation without a diffusion

Fig. 5. Cell density of filamentous (solid lines) and non-
filamentous (dashed lines) species in two cases: d and e (refer
to the text).
process. In this verification, the time for the 30-day
simulation was  2 h using PC 586/550 MHz.
The results indicate that the non-filamentous
biomass transitions of the two cases have a similar
pattern, differing only by  10% (Fig. 5). Although
the transport is extremely hampered inside the
viscous sublayer, nearly a sufficient amount of
nutrients is supplied for periphyton growth at a
diffusion coefficient of 5× 10 − 9 m2 s − 1.
However, a temporal variation of the internal
nutrient concentration distinguished cases (d–e)
(Fig. 6). With Kdiff = 5×10 − 9 m2 s − 1 (Fig. 6, case
(e)), the nutrient flux is not sufficient to rapidly
increase the filamentous algal biomass from day 12,
thus showing a reduction in the internal nutrient
concentration. A similarity between Fig. 3(case (a))
and Fig. 6(case (d)) implies that at a diffusion
coefficient of 1×10 − 6 m2 s − 1, the rate of nutrient
flux is sufficient for each cell regardless of the
community density.
Fig. 7 shows marked differences in the nutrient
resource indices at different diffusion coefficients in

Fig. 6. Temporal variation of the internal phosphorous con-

centration of filamentous (solid lines) and non-filamentous
(dashed lines) cell in the bottom layer in two cases: d and e
(refer to the text).

fluctuation of a filamentous community causes

fluctuations in the light resource index of both
species (Fig. 4, cases (a – b)). A comparison of Figs.
2 and 4 implies an exact inverse correlation between
the filamentous algal biomass and the light resource
condition.

4.2. Varied external nutrient concentration

The second verification was made with two

values of the diffusion coefficient inside the viscous
sublayer: (d) 1×10 − 6 m2 s − 1, which was in the
same order with the molecular viscosity; and (e)
5× 10 − 9 m2 s − 1, which is comparable with the
molecular diffusion coefficient for oxygen of 2×
Fig. 7. Temporal variation of the phosphorous (solid lines),
10 − 9 m2 s − 1 (Sand-Jensen and Revsbech, 1987).
nitrogen (dashed lines), and light (dotted lines) resource index
Eq. (13) was used to calculate the diffusion coeffi- of filamentous (left) and non-filamentous (right) cell in two
cient outside the viscous sublayer. The time step cases: d and e (refer to the text). Arrows indicate detachment
was 5 s, which was much smaller than the value of filaments.
70 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75
Fig. 8. Vetical profile of velocity on day 5 (d5), day 15 (d15),
and day 25 (d25).
the viscous sublayer. The phosphorous resource
index of filamentous species was stable at  0.5
with Kdiff = 1×10 − 6 m2 s − 1, exactly the same
as with the infinite diffusion coefficient, while it
gradually decreased to 0.4 on day 18 with
Kdiff = 5×10 − 9 m2 s − 1. This difference, how-
ever, did not affect much the biomass (Fig. 5).
The similarity between cases (a) and (d) in the
biomass (Figs. 2 and 5), the internal nutrient
(Figs. 3 and 6), and the resource indices (Figs. 4
and 7) indicates that the diffusion coefficient of
1× 10 − 6 m2 s − 1 is still high enough to support
the dense community. As the nutrient require-
ment of filamentous species was higher than
non-filamentous species (Table 1), the nutrient
resource index of filamentous species was always
smaller in both cases than non-filamentous spe-
cies.
Fig. 8 shows the vertical velocity distributions
on days 5, 15, and 25. The periphyton commu-
nity built on short turfs of filaments markedly
modifies the vertical profile of velocity such that
it de-creased near the bottom, but compensated
for this by increasing above the middle of the
depth on day 15. With further increasing
biomass as much as to form a mat on the sur-
face (on day 25), however, the surface velocity
decreases and the depth of the maximum value
reduces until at about two-thirds of the total
velocity (38 mm). The final velocity profile
agrees generally with the experimental observa-
tion that most of the water flow occurs in the
middle of the floating mat of zygnematalean and
the bottom (Peterson and Stevenson, 1990).
5. Application
Observations made at a stream facility at the
University of Louisville, Kentucky, USA
(Peterson and Stevenson, 1990, 1992) were used to
validate the model. To provide a water velocity of
0.29 and 0.12 m s − 1, water depths of 2.5 and 6.0
cm, respectively, were used. Despite fluctuations
were expected in the experiments, phosphorous
and nitrogen concentrations were kept constant at
the entrance of the channels throughout the
simulation at 2.0 and 15.0 mg l − 1, respectively, and
the water temperature was 17.8°C.
Fig. 9 shows the simulated results on cell
densities of these current regimes compared with
observations. Generally, a community subject to a
high flow rate experienced a markedly smaller
increasing rate and a lower peak biomass than a
community subject to a low flow rate. In the slow
current regime, the peak biomass of A.
minutissima was 9500 cell mm − 2 on day 6 and
again occurred on day 14 with a higher value of
12 000 cell mm − 2, while in the fast current regime,
it was only 3200 cell mm − 2 on both days 15 and
30 due to the lower initial density and the higher
net emigration rate.
The Spirogyra spp. biomass started dislodging
on day 25 with its high biomass of  15 000 cell
mm − 2 and fluctuated in the last 10 days of the
experiment in the slow current regime, while in the
fast current regime the dislodgment was delayed
until day 30 with 4000 cell mm − 2. The lower
biomass of Spirogira spp. in the fast current
regime was mainly because of the lower
immigration rate than in the slow current regime.
Due to the low biomass, the time of dislodgment
was delayed 5 days in the fast current regime.
Fig. 10 shows the variations in the resource
index of three species at the bottom. Having a high
light saturation level, Spirogyra growth was
always subject to the light limitation of both
current systems. Although slightly higher in the
fast current due to the lower biomass density, the
light resource index of Spirogyra was maintained
at  0.2 until its biomass increased and shaded
itself. In the later stage of the experiment, how-
ever, all species suffered from light limitation near
 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75
the bottom, which was again more serious in the
slow current regime.
From the initial values, the nutrient indices of
algae tended to decrease because of the high algal
consumption of nutrients under sufficient irradia-
tion. After Spirogyra arrived at the water surface,
the high floating biomass interrupted the penetra-
tion of solar irradiation into the water column,
and thus the nutrient concentration gradually re-
covered at the bottom layers. As a result, in the
final stage of growth, nutrient resources were
nearly satisfactory for all species shifted from
having nutrient-limitation to light-limitation. In
both flows, the light resource index of all species
temporarily fluctuated because of the dislodge-
ment of Spirogyra at the later stage of the
experiments.
Because Spirogyra has a high nutrient require-
ment, its internal nutrient concentration varied
largely according to the ambient concentration
compared with that in diatom cells, especially in
the slow current. For A. minutissima and Synedra
spp., however, the ambient nutrient concentra-
tions were relatively sufficient and thus the nutri-
ent resource indices of these species were stable at
Fig. 9. Comparison of cell density between computed (lines) and observed (symbols) results in the slow flow (left) and fast flow
(right) regime. SP, Spirogyra; AM, Achnanthes minutissima; SYN, Synedra spp.; Dead, Dead Achnanthes minutissima.
71
0.5 –0.8 and 0.6 –0.8, respectively, in both current
regimes.
After reaching the water surface, Spirogyra
filaments started to extend along the stream. An
increase in density of filaments near the surface
reduced markedly the external nutrients in both
current regimes (Fig. 11). In the surface layers,
phosphorous and nitrogen concentrations were
reduced from the initial values to 1.2 and 2.0 mg
l − 1 in the slow current on day 24 and to 1.8 and
10.0 mg l − 1 in the fast current on day 28. With a
high Spirogyra biomass in the final stage, the
nutrient concentrations in the water fluctuated
with the frequent detachment of algal community.
At the bottom, the external nutrient concentra-
tion decreased with the increasing biomass until
0.5 mg l − 1 in phosphorous and 1.5 mg l − 1 in
nitrogen on day 15 in the slow current and only
until 1.0 and 3.5 mg l − 1 on day 30 in the fast
current. In the slow current regime, however,
nutrients at the bottom quickly recovered after
the floating filamentous mat formed on day 15,
because the growth rate and the nutrient uptakes
of diatom species declined.
72 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75

Fig. 10. Resource indices of three algal species at the bottom layer in the slow flow (left) and fast flow (right) regime: P, Phophorous;
N, Nitrogen; L, Light.

The benthic algal biomass temporally varied
with successive rate, autogenic dislodgment, and
disturbance of the surrounding environment. Both
physiological and morphological characteristics
determine the competitiveness of algal species in
the benthic mat. Competitive dominants under
favorable environmental conditions are species
that have the highest intrinsic increasing rates to
allow them to proliferate over other taxa. Despite
prominent differences in the nutrient concentra-
tion between the bottom layer and the overlying
water and between the fast and slow currents,
nutrient resource index of the diatom, A. minutis-
sima and Synedra, was relatively stable. In other
words, A. minutissima appeared to have adapted
to exploit relatively low levels of dissolved nutri-
ents. This agreed with the findings of Pringle
(1990).
A Spirogyra recession occurring at the end of
both currents was reproduced fairly well by this
study, implying that light availability to cells in
the lower layers of periphyton matrices regulated
biomass accumulation by affecting the strength of
attachment of the bottom cells to the substratum
(Boston and Hill 1991; Peterson, 1996). Several
pieces of evidence suggest that Spirogyra is not
capable of tolerating prolonged exposure to ex-
tremely low or no light (Graham et al. 1995).
 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75
Fluctuations of diatom biomass and A. minutis-
sima dead cells coincided with the Spirogyra re-
cession due to the secondary detachment of
non-filamentous cells inhabiting the interstices of
the filamentous mat. Also, it shows that, a consid-
erable number of dead cells were settled from the
overlying water and were trapped on top of the
benthic mat and then detached with filaments.
6. Concluding remarks
This study numerically indicated the impor-
tance of the diffusion process in nutrient exchange
between the periphyton community and overflow-
ing water. The model showed reasonable behavior
of periphyton communities, such as light reduc-
tion in the periphyton mat, variation of the inter-
nal nutrient concentration, and the transition
process of a vertical velocity profile.
As several effects were minimized in this study,
such as the respiration effect on the cell cytoplasm
volume, nutrient releases from the sediment, or
biological acclimation of the benthic algae to re-
Fig. 11. Temporal variation of nutrient concentration in the slow flow (left) and fast flow (right) regime.
73
sponses of resource-limited environments, erosion
of the viscous sublayer due to the existence of the
biomass, etc. this model should be applied with
some cautions. In the model of this study,
biomass loss due to grazing was simply treated as
a constant rate though it should be varied with
herbivorous species and density. The fact is worth
emphasizing that taxonomic and physiognomic
structures of periphyton community species are
markedly affected by the presence of invertebrates
(Mulholland et al., 1994). All these factors being
taken for granted, however, the model is consider-
ably useful to study the response of a periphyton
community to a particular environmental factor.
Finally, to deal with the diffusion process of a
high biomass concentration, a time step of min-
utes or even smaller should be used.
Acknowledgements
The study was made while the second author,
Duong Hong Son, was receiving a scholarship
from the Japanese Ministry of Education, Culture
74 T. Asaeda, D. Hong Son / Ecological Modelling 137 (2001) 61–75

and Sports. This study was financially supported Detachment condition for the bottom-attached
by the Japanese Ministry of Education, Culture, filaments:

) )
and Sports, Foundation of River and Watershed
Management, and the Maeda Engineering Foun- Nn, k, i Dt − 1
dation. These supports are gratefully acknowl- Ndetach = if: zCdU 2Si Ln, i
0
edged. We also acknowledge V.T. Ca for his
valuable comments on the flow dynamics part of
the study. ] RFcr1
(6A)
B RFcr1

Appendix A
Detachment condition for the laterally attached
filaments:

F F
The following formulae are used in the model.
Incline angle of filaments, h:
Buoyancy (z −za )gVi KzV 1/3
tan h :
i
= = where
Ndetach = Nn, k, i Dt
−1
Drag zCdU 2V 2/3
i U2 if: zCdU 2Si Ln, i
0
(z −za )g
Kz = (1A)
zCd B
] F
DV cr2
where: Periphyton settlement:
 
(7A)
B
k−1 B F
DV cr2
n, k, i( j ) = max 0, Cn, 0, i( j ) − % C n, kk, i( j )
C initial initial

kk = 1

Bk
S Dz
n (2A)
Detachment rate non-filamentous community:
DVk nor
Fdetach
The light attenuation constant and light resource Gdetachj = G0 where
F0 + Fdetach
index:
zCdU 2Sj
Bk Fdetach = (8A)
pk = pw +pc and B/DV
DVk

IRn, k, i( j ) =
Ik 
exp 1 −
Ik  (3A)
Isati( j ) Isati( j )
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