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CICEET progress report for period 08/01/2000
through 01/31/2001
Project Title:
Modeling the effects of changes in turbidity on light available
for submerged aquatic vegetation
Project Coordinator:
Roger I. E. Newell
Professor, Horn Point Laboratory, University of Maryland Center
for Environmental Science
PO Box 775, Cambridge, MD 21613
Phone: 410-221-8410, Fax:410-221-8490
Email: newell@hpl.umces.edu
Additional Principal Investigators:
Raleigh R. Hood
Assistant Professor, Horn Point Laboratory, University of Maryland
Center for Environmental Science
PO Box 775, Cambridge, MD 21613
Phone:410-221-8434, Fax:410-221-8490
Email: raleigh@hpl.umces.edu
Evamaria W. Koch
Assistant Professor, Horn Point Laboratory, University of Maryland
Center for Environmental Science
PO Box 775, Cambridge, MD 21613
Phone:410-221-8418, Fax:410-221-8490
Email: koch@hpl.umces.edu
Raymond E. Grizzle
Associate Professor
Jackson Estuarine Laboratory
85 Adams Point Road
Durham, NH 03824
Phone: 603-862-2175, Fax: 603-862-1101
I. Accomplishments
A. Scheduled Tasks
Undertake in October 2000 the second of the two proposed field
studies to quantify sediment resuspension in SAV beds when,
due to normal senescence, the seagrass shoot density will be
lower than during the June study period.
Complete the experimental work in the laboratory to measure
changes in light extinction coefficients associated with oyster
and clam feeding over a range of seston concentrations at 15,
20, and 25oC water temperatures.
Continue the development of our mathematical model in STELLA.
Our first goal is to predict, on theoretical grounds, the interactions
between seagrasses and suspension feeders through their effects
upon suspended sediment concentrations and light penetration.
Start to parameterize the model using our own field and laboratory
data.
B. Progress on Tasks.
1. The influence of seagrasses on sediment resuspension has
been measured in June and October in Duck Point Cove at Bishops
Head Point at the entrance to Monie Bay, Maryland. In this same
cove we selected two sites, one densely vegetated by the seagrass,
Ruppia maritima, and one unvegetated. At each site, two small
platforms were mounted to hold the necessary equipment to monitor
environmental conditions in a vegetated and an unvegetated area
over two 10-d periods (Summer when the vegetation occupied the
entire water column and Fall when the plants only occupied a
small fraction of the water column). We used an ISCO automated
water sampler at each site to collect 1 l of water every 2 h
to determine total suspended solids (TSS), grain size distribution,
and salinity. Additional abiotic parameters recorded included
water temperature, light availability (4 LiCor sensors)
and wave characteristics. At the end of the experiment, seagrass
density, canopy height, and photosynthesis versus irradiance
curves were measured. Samples are in the final stages of being
analyzed. Additionally, the critical erosional threshold of
the sediments at each site was quantified using microcosms under
controlled conditions
2. We used adult oysters (Crassostrea virginica) (8 to 10
cm shell height) collected from the Sandy Hill oyster bar in
the Choptank River. Hard clams, Mercenaria mercenaria ,(shell
height of 5 to 6 cm) were grown in Plantation Creeks, VA and
provided by Dr. Mike Pierson of Cherrystone Aquafarms. Both
species of bivalve were collected in March and August 2000 and
held in the lab for 14 to 20 d acclimation prior to use in the
feeding studies. In March one group of each species was held
at the field ambient water temperature of 15oC and
one group was acclimated to 20oC.
In August animals were maintained only at the field ambient
water temperature of 25oC. Oysters were held in flowing
ambient estuarine water salinity (salinity 12 to 15 ppt) with
a natural seston complement. Clams were held in non-flow through
aerated tanks with 20 ppt water made by mixing estuarine water
with full strength seawater with 20% of the water exchanged
every second day. Clams were fed algal paste (Chgra) at 2% of
dry body weight per day as a maintenance ration. Clams were
put into 12 cm deep plastic beakers containing coarse sand into
which the deeply buried during the acclimation period.
The influence of eastern oysters and hard clams on turbidity
and light penetration was evaluated in 1 m deep 1000 l tanks
filled with estuarine water. These tanks have a continuous mixing
systems consisting of rotating, reversing paddles, with speed,
direction, and duration controlled by computer to simulate mixing
in nature. This system ensures homogeneous mixing of the water
column without resuspending sediment from the bottom as described
by Sanford (1997). Light was provided by 4 ft high output flourescent
tubes which produced irradiance of ca 200 ( µmol photons m-2
s-1) just below the water surface
In separate experiments oysters and clams were placed on the
bottom of the tanks and allowed to feed undisturbed for 2 h.
Oysters were placed in 2 cm deep plastic pots to facilitate
retention of both feces and pseudofeces and clams were placed
directly in the tank while still buried in the sand-filled plastic
beakers. At intervals for between 10 and 24 h the diffuse attenuation
coefficient for photosynthetically available light, Kpar
was measured using a 4 Li-Cor light meter positioned
just beneath the water surface (O) and at 0.5 m (Z) beneath
the surface. The flourescent lights were only switched on for
the 10 min period required to make readings and were then turned
off to reduce any phytoplankton growth during the experimental
runs. Due to slight variations in light output from the high
intensity flourescent tubes we measured light levels (µmol photons
m-2 s-1) haphazardly 4 times in each of
the 3 tanks. The 4 light readings were then used to calculate
the extinction coefficient
Kd = [ln (light Z /Light O )] / Z from which an
average Kd was calculated.
In order to estimate the loss in light intensity associated
with distance from the light source, rather than attenuation
by material in the water, we filled these same tanks with tap
water containing no suspended particles and, as before, measured
light just under the water surface and 0.5 m further into the
tank. This Kd value was used to correct all of the
experimental Kd values. Regression equations of Kd
against time were calculated for each data set and used to calculate
the absolute change in Kd (increase in light penetration) for
a 24 h period. The change in Kd for the control tank associated
with particle settlement was then subtracted away from the experimental
values.
Water was collected concurrently with the light measurements
using a siphon hose close to the middle of the tank. Particle
size distribution and abundance of particles > 2 µm (size
above which clams and oysters can retain a large percentage
of suspended particles) was counted on subsamples using a Coulter
Multisizer II. Water samples were filtered through 47 mm Whatman
GF/C filters for analysis of seston dry weight (mg l-1)
in order to quantify the relationship between seston and Kpar.
Filters were rinsed with isotonic Ammonium formate to remove
salt (Armstrong 1958, Berg and Newell 1984). Changes in chlorophyll
concentration due to bivalve feeding was measured at each water
collection by filtering water and extracted for chlorophyll
a analysis (µg l-1). At the end of each experiment,
oyster and clam tissues were individually dissected from shells
into pre weighed aluminum pans and dried at 80 oC
for 2 d for determination of each animals total dry tissue
weight.
Aggregate clearance rates for the known biomass of bivalves
in each tank (liters water cleared h-1 g-1)
were calculated based on the rate of particle disappearance
in tanks. Particle concentrations measured using the Coulter
Multisizer at intervals in each tank were transformed by logarithm
and used to prepare a linear regression of particle disappearance
over time for each tank. These equations were used to predict
the initial (time 0; Ci) and final (time 24 h; Cf)
log transformed particle concentrations in each tank, including
control tanks without bivalves (time 0; Cci and time
24 h; Ccf) in order to correct for particle settlement
during the feeding period.
These values were then inserted into the following equation
(Coughlan 1969) to calculate aggregate clearance rate (CR):
CR (L h-1) ={ [ (loge Ci-
loge Cf)]- [(loge Cci
- loge Ccf)]} x (V/t)
where V = Tank volume (1000 L) and t = duration of feeding
period. Clearance rates were corrected to a dry tissue weight
of 1 g by dividing by the total dry tissue weight of all animals
placed in that tank.
3. We have continued to make substantial progress in the development
of our mathematical model predicting the interrelationships
between seagrasses and suspension feeders on light penetration.
C. Difficulties Encountered
No major difficulties have been encountered so far in the
development of this project.
D. Anticipated Success in Meeting Project Objectives in Scheduled
Project Period.
We have met all the objectives we set out for the first 18
months of this 24 month project
E. Preliminary data
The field studies in the seagrass beds and adjacent unvegetated
areas are showing the contribution of SAV to decreasing total
suspended solids (TSS) loads in the water column. This is a
process driven by physical processes (waves) as well as the
reproductive state of the plants. The concentration of TSS increases
with increasing wave height in both the vegetated and unvegetated
areas (Figs 1 and 2).
Attenuation of waves is a function of the amount of the water
column that is occupied by the vegetation. In the Summer, when
plants are in a reproductive state, and hence occupy the entire
water column, wave attenuation is high. In the Fall, when plants
are in a smaller vegetative state, and hence occupy only a small
fraction of the water column, wave attenuation is low (Fig.
3). Based on this we might expect that the overall TSS levels
within the beds would be higher in the Fall than in the Summer.
Our data suggest that in fact the highest loads are in the Summer
most likely due to the high contribution of particulate organic
matter from phytoplankton during the warmer months. In addition,
because there are more seagrass leaves in the summer there is
a greater area upon which material from the water column can
settle and later be resuspended. Independent of the source of
suspended particles, light attenuation in the water column increases
with increasing TSS concentrations (Fig. 4).
Figure 1. Wave dependent resuspension of particles in a vegetated
(black symbols) and adjacent unvegetated (white symbols) area
when the vegetation occupied the entire water column (Summer).
Figure 2. Wave dependent resuspension of particles in a vegetated
(black symbols) and adjacent unvegetated (white symbols) area
when the vegetation occupied only a small portion of the water
column (Fall).
Figure 3. Wave attenuation of Ruppia maritima when reproductive
(shoots occupy the entire water column black symbols)
and when in its vegetative state (shoots occupying only a portion
of the water column white symbols).
Figure 4. Light attenuation (Kd) as a function of total suspended
solids concentration (TSS) in a SAV habitat.

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2. The data from the completed oyster and clam feeding studies
(Table 1) show that there were clear and measurable changes
in the light attenuation coefficient (Kd) over the
course of the feeding period. Our study provides the first quantification
of how the removal of suspended particles by bivalves increases
the amount of light that can penetrate through the water column,
with rates being greatest for both species at 25oC.
As anticipated, the weight-specific feeding activity of oysters
was much greater than the clams. This is due to the fact that
oysters are better adapted to maintaining high rates of particle
clearance in turbid waters. The oyster's feeding strategy relies
on the highly efficient sorting capacity of their gills and
labial palps that allows them to reject non-nutritious and excess
particles as pseudofeces and hence ingest a constant amount
of nutritious particles. In contrast, clams produce fewer pseudofeces
and regulate ingestion by reducing clearance rates. These data
will then be used to parameterize the oyster and clam filtration
component in the simulation model described below.
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F. Continued Model Development:
We have essentially completed the formulation and parameterization
of the predictive model and implemented it in Stella. As described
in our previous reports, the basic model is composed of two
equations, one which describes the rate of change in SAV biomass
(Bsav):
1) dBsav /dt = um(1-e^(-Iavg/Ik))Bsav
- rBsav,
And a second which describes the rate of change of suspended
seston concentration (S):
2) dS/dt = [M(b-c)
+ wsS B CbSBb]Zbot
In (1) um is the maximum growth rate of SAV, Iavg
is the average irradiance over the length of the shoot, Ik
is the light saturation parameter for SAV photosynthesis and
r is coefficient characterizing respiratory losses. In (2) M
is the erosion rate, b and c
are the bottom shear stress and the critical shear stress for
resuspension, respectively. In addition, in (2) ws
is the sinking rate of seston, Cb is the filtration
rate of bivalves, and Bb is the biomass of bivalves
(the latter is specified, i.e., not dynamically modeled). Zbot
is the distance from the water surface to the bottom. The model
is cast in a vertically integrated form and with spatial units
of m2. Equation (2) feeds back on (1) through S,
which partly determines Iavg in (1). S is determined
in 2) by the balance between sedimentation, bivalve filtering,
and sediment resuspension.
In order to calculate Iavg we include light attenuation
due to water and dissolved substances, Kx, seston,
Ks, and self-shading by SAV, Ksav. And
we split this attenuation into two parts, attenuation above
the shoots, K1, which is due only to water, dissolved
substances and seston:
3) K1 = Kx + Ks,
and attenuation below the tips of the shoots, K2,
which include all of the above plus self-shading:
4) K2 = Kx + Ks +
Ksav.
For simplicity we assume that Kx is a constant.
We estimate Ks = m1S from seston concentration
as described above, and we estimate Ksav = asavBsav
using an SAV biomass specific attenuation coefficient,
asav. The incorporation of a self-shading effect
in the model allows us to dynamically model maximum bed density
as a function of depth and light attenuation in the overlying
water column.
The average light over the shoot length is determined by first
calculating the irradiance at the top of the shoots, Ztop,
where light is attenuated by Kx and Ks:
5) Itop = Ioe-K1Ztop
and then averaging the light from Ztop to the bottom,
Zbot:
6) Iavg = Ioe-K1Ztop(e-K2Ztop
- e-K2Zbot)/[K2(Zbot
B Ztop)]
where Io is the irradiance at the water surface.
For this application we assume that the critical bottom stress
for resuspension, ôc, is a constant, and that
the bottom stress, ôb is determined primarily
by waves:
7) b = Hrñ(õ(2_/T)3)1/2[2sinh(2_Zbot/L)]-1
In 7) Hr is the wave height, ñ is the density
of the water, õ is the kinematic viscosity of the water,
T is the wave period, and L is the wavelength. The relationship
between wavelength and period is calculated using standard shallow
water formulae.
Equation 7 does not include wave damping effects due to the
presence of SAV. Yet a key aspect of this model is the specification
of these effects, i.e., how do changes in the density and height
of SAV modify waves and how does this, in turn, change the bottom
stress, ôb. To our knowledge, there are no
published quantitative relationships that describe these effects
for SAV beds in shallow water environments. We are currently
working on deriving relationships that can be used to modify
Hr as a function of bed density and SAV height using
observations collected by Koch as part of another study.
The model has been implemented in STELLA and preliminary runs
(with Ztop = 0 and ôb = 0, and the
SAV model parameterized to simulate Rupia) give realistic depth
distributions, i.e., equilibrium SAV bed densities declining
exponentially from about 3000 shoots m-2 at Ztop
= 0 m, to 0 shoots m-2 at Ztop = 5 m depth.
II. Tasks and activities for next reporting period
A. Tasks for the next reporting period
- Formulate and implement a relationship which describes wave
height as a function of SAV height and bed density.
- Complete the development of the predictive mathematical model
incorporating all functional relationships and parameters derived
from the field and laboratory studies. Carry out a full sensitivity
analysis on the model.
- Characterize the model-generated interactions between wave
height, SAV growth and bivalve feeding.
- Test the predictions of light penetration and seagrass growth
from our STELLA model at clam aquaculture farms on the lower
eastern shore of Chesapeake Bay where turbidities may have been
reduced sufficiently to allow seagrass beds to become reestablished.
- Develop the user interface of the model. Make verbal presentations
at scientific meetings and hands-on demonstrations of the model
to resource managers. Make any needed improvements and refine
the user interface.
- Distribute the model via the internet.
- Write paper describing the model and the results of the field
research in scientific journals.
B. Work plan to accomplish tasks
We will complete the development and testing of our mathematical
model in STELLA. The model will incorporate the data from the
field studies of the seagrass beds at Bishops head and the laboratory
bivalve feeding studies described above. Our first goal will
be to predict the interactions between seagrasses and suspension
feeders through their effects upon suspended sediment concentrations
and light penetration. This model will be completed before we
start our second summer of field work in May 2001.
We will test our model predictions of the influence of bivalve
feeding on light penetration and seagrass growth at Cherrystone
hard clam aquaculture cooperative on the lower eastern shore
of Virginia on the Chesapeake Bay. We have been in contact with
the manager, Dr. Mike Pierson, and have selected two clam grow-out
sites with different sediment characteristics and exposure to
wave action. Mattawoman creek has quite a sandy bottom and is
exposed to NW waves from Chesapeake Bay. In the upper part of
Cherrystone inlet clams are grown in a sheltered location with
a muddy bottom.
We have made a strong start in disseminating the results of
this work to the management and research community. We have
been invited to make the following scientific presentations
concerning our research:
Newell, R.I.E. Potential for N, P, and Sediment Removal by
Oysters. Invited presentation at an AExploratory meeting on
nutrient/sediment removal by oysters@ organized by EPA Chesapeake
Bay Program. Annapolis MD March, 2001
Newell, R.I.E., E. Koch and R.R. Hood. Modeling influence
of populations of suspension-feeding bivalves and seagrasses
on suspended particulate load and consequent changes in light
attenuation. Invited presentation Estuarine Research Federation
Biennial Meeting. November2001.
C. Concerns or difficulties
At this point we anticipate no major difficulties regarding
the future progress of this project.
III. Expenditures
In no budget categories are expenditures exceeding estimates.
Apparently invoicing from the University of Maryland to the University
of New Hampshire is behind schedule. Our business office realizes
the problem and told me that they would take care of it immediately.
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