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Structure and Functioning of a Managed Grassland Ecosystem
CONTENTS
SUMMARY
An area of seeded grass has been subject to two management regimes for
more than 20 years. One area has been mown at intervals of two to three
weeks throughout the growing season, while an immediately adjacent area
has been left uncut and has undergone successional development and has
now developed scrub cover (see photograph). The effects of mowing on the
structure and functioning of the grassland are investigated by comparing
mown and unmown plots.
The results demonstrate:
- how management selects for plants with particular adaptive traits
- interactive effects of management on the whole biological structure
and functioning of the grassland system
- biophysical controls on carbon and nitrogen dynamics
- inputs and losses of nutrients by rainfall and leaching of soil solutions
- effects of vegetation cover on heat fluxes and nitrogen dynamics.
The experimental area

The site in 1998. The unmown area on the right reflects about
20 years of successional development, with some scrub encroachment
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KEY WORDS
grassland, ecosystem analysis, carbon, nitrogen, succession, mowing,
plant diversity, invertebrate diversity
AUTHOR
Corresponding author: |
Professor Jonathan M. Anderson |
Address: |
School of Biological and Chemical Sciences, Hatherly Laboratories,
University of Exeter, Exeter EX4 4PS, UK |
Email: |
j.m.anderson@exeter.ac.uk
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INTRODUCTION
Aim
To investigate the effects of mowing on grassland vegetation diversity
and succession, and on nutrient pools and fluxes in plants and soil.
Division of the project into separate exercises
- Because of the large and comprehensive nature of this project it
has been divided into two parts. Part I is an initial survey of plant
diversity (herbs and shrubs) in mown and unmown plots, a study of adaptive
traits of plants to intensive mowing and a comparison of enumeration
methods. Part II is a system analysis comprising thirteen separate exercises
(A to M), six related to pools and seven to fluxes. These exercises
are described individually.
- The ecosystem project is best tackled in its entirety. The exercises
are carried out singly or in combinations by separate small groups of
students. Some exercises are more time-consuming than others, so shorter
ones can be combined in single practical sessions.
- Alternatively, if resources are limited or a less comprehensive approach
is required, the project may be reduced to a single exercise or a combination
of a few of the exercises. Obviously, the value of the original holistic
approach will then be greatly diminished.
- The results of the individual exercises are synthesised at the conclusion
of the investigations. If all the exercises have been tackled an overall
picture of the linked processes occurring within the grassland system
and the differences caused by the two management treatments will be
apparent.
- This project outline is not a comprehensive methods schedule, since
background knowledge and technical backup is required for many of the
exercises. The exercises can be used separately and/or be modified as
required. The illustrative data for each exercise are from student practicals
and may not necessarily show expected trends. The protocol can, however,
be refined for research into ecosystem processes.
Table 1 is a summary of the complete system project components.
POOLS
| EXERCISE |
COMPONENT |
PARAMETERS |
METHODS |
| A |
Above ground vegetation |
Living and dead material (g m-2) |
Clipping and weighing material from quadrats |
| B |
Below ground vegetation |
Roots and stolons (g m-2 at 0-5
cm, 5-10 cm depth) |
Core sampling and washing through sieve |
| C |
Soil organic matter and moisture |
Soil bulk density (g cm-3). |
Core sampling, wet and dry weights |
| Organic carbon (% by weight) in soil and in
above ground vegetation. |
Loss on ignition (LOI) |
| D |
Nutrient pool: plants |
Na, K, Ca and N in above and below ground
vegetation (mg g- 1) |
Wet digestion, grinding, element analyses |
| E |
Nutrient pool: soil |
Exchangeable Na, K and Ca in soils. |
Extraction by ammonium acetate, element analyses. |
| Total N in soil. |
Wet digestion, element analysis. |
| F |
Fauna: earthworms |
Number and wet mass (g m-2) |
Soil sorting |
| surface-living macrofauna |
Number of individuals per taxon |
Pitfall traps |
| soil fauna |
Number of individuals per taxon |
Extracted core samples |
FLUXES
| EXERCISE |
COMPONENT |
PARAMETERS |
METHODS |
| G |
Heat |
Temperature |
Max/min thermometers |
| H |
Precipitation |
Rainfall (water, nutrients, pH) |
Rainfall collectors, rainfall measurement, element analysis,
pH metered |
| I |
Soil water |
Infiltration (ml h-1) |
Ring infiltrometers |
| J |
Carbon fluxes - CO2 |
'Soil respiration' |
CO2 absorbed in alkali trap in field and
laboratory |
| K |
Nitrogen mineralisation |
NH4+ and NO3-
mineralisation |
Soil incubation in field and laboratory |
| L |
Denitrification |
N2O production (in open and under tiles) |
Determination by gas chromatography |
| M |
Soil solution sampling |
Mineral-N and SO4-- |
Porous cup samplers, element analysis |
| pH |
pH metered |
Learning objectives
- Understand how to carry out an ecosystem study.
- Appreciate group objectives and team responsibilities.
- Understand experimental design, sampling methods and the importance
of replication.
- Gain practical experience of a wide range of field and laboratory
methods used in ecological studies.
- Understand processes of nutrient cycling between plants, invertebrates,
litter and soil.
- Gain experience of the identification of herbs and diverse invertebrate
groups to practical taxon (at least Class/Order).
- Understand the impacts of meadow management (mowing) on
- plant diversity
- plant succession
- grass growth, biomass and nutrient quality
- nutrient pools and fluxes
- microclimate and habitat structure
- surface-living and soil invertebrate communities.
- Understand the role of earthworms in soil structure and hydrology.
Ecological concepts
Vegetation management has fundamental and interactive effects on the
structure and functioning of ecological systems. Different intensities
of management (mowing) affect the composition of plant and animal communities
and this has implications for different conservation objectives for flowers,
butterflies, birds, mammals, etc.
Skills required
A level Biology and/or Geography
GCSE/AS Mathematics and Chemistry
Level of study
Undergraduate year 1 (for exercises on plant and invertebrate communities)
or year 2/3 (for exercises on system analysis).
LEGISLATIVE AND ETHICAL CONSIDERATIONS
Health and safety
- The use of chemicals in the laboratory must be subject to COSHH regulations.
- Documentation available in the laboratory advises students of risks
arising from their work and precautions to be taken where necessary.
- Precautions to be taken in the laboratory and with the use of reagents
are mentioned in the Methods sections. Care should be taken with barium
chloride (moderately poisonous). This could be omitted from CO2
procedures, but with loss of analytical precision.
Disability discrimination
All the exercises could be carried out within the constraints of particular
disabilities and accessibility of the field sites.
Animal welfare
- Extraction of earthworms from soil has traditionally been done using
0.25% formaldehyde solution, but its general use is now prohibited .
Mustard and chilli solutions have been used to expel worms from soil,
followed by washing to remove the irritants. However, digging up and
hand sorting soil samples, although more laborious, is preferable, not
only because it harms few earthworms but it is more effective for sampling
juveniles and certain soil-living species. The process of hand-sorting
also provides insight into the nature and structure of soil.
- Pitfalls containing a trapping solution inevitably result in the
death of invertebrates (and very occasionally small mammals). It is
impractical to use dry cups. Extraction of soil invertebrates into preservative
kills thousands of small animals, but this is rarely regarded as a matter
of concern by students, and raises interesting ethical issues about
animal type and size.
Environmental awareness
The students should appreciate the value of long-established experimental
plots and carry out their investigations with the minimum of disruption
to the site. If the field area is small it is advisable to fill in holes
using soil from the site and to avoid using the same plots the following
year.
METHODS
Staffing and student group size
30-60 students in about a dozen small groups require at least 3-4 well-briefed
demonstrators for the system analysis, because of the range of different
exercises involved. The main recurrent problems are inadequate labelling
of samples, and bulking materials rather than keeping separate replicates
as a measure of sample variability.
Study site
An area of seeded grass on the University of Exeter Estate is used for
this project. For more than 20 years half of this experimental area has
been mown (without the mowings being removed) and half has been left unmown
(Figure 1).
Figure 1. Sketch of the site layout

Time of year
April to October
Data handling and statistical techniques
Where appropriate the following are used for analysis of data gathered
during the exercises:
- Spread sheets
- Regression (calibration of readings and standards, or interpolate
graphically).
- ANOVA (comparison within and between treatments blocks if sufficient
sampling intensity)
- t-tests (comparison of means).
Preliminary field procedure
A long-term experimental site is set up in grassland.
- One area is mowed at intervals of at least 2-3 weeks throughout the
growing season, with the mowings not removed, while an immediately adjacent
area is left uncut.
- The methods described here could be used on any paired treatments.
These need not necessarily be contiguous, but the soils and other environmental
conditions should be identical as far as possible, and have a record
of past management.
- This is a long-term experiment, but sampling can commence one year
after the experimental site has been set up. Selective effects of mowing
on plant species can be observed within one season.
- One 10 m by 20 m plot is set up bridging the mown and unmown grassland
areas (Figure 1). Alternating 1m rows are used for sampling and as paths.
Students must avoid trampling the area until the plots are established.
- Replicate samples of plant material and soil are taken from mown
and unmown plots, as detailed in the separate exercises.
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