Академический Документы
Профессиональный Документы
Культура Документы
Hawes, C., Begg, G. S., Squire, G. R. and Iannetta, P. P. M. 2005. Individuals as the
basic accounting unit in studies of ecosystem function: functional diversity in
shepherds purse, Capsella. / Oikos 109: 521 /534.
Agricultural intensification has led to a systematic erosion of the biodiversity in arable
ecosystems. Despite this, densities of plant species in the seedbank of arable fields are
still sufficient to provide the potential for recovery of the arable flora and associated
fauna. Identification of management practices to achieve this, while minimising
negative effects of weed competition on crop yield, requires a mechanistic
understanding of functional diversity in arable systems. However, a review of the
ecological, physiological and genetic characteristics of 66 representative UK arable
weed species revealed major gaps in this knowledge for even the most common species.
Even less is known about the degree of variation between individuals within these
species that contributes to overall levels of functional diversity. Classification of
organisms into functional groups on the basis of species-level taxonomy is inadequate
to describe the functional diversity of a system, since variability in a particular
physiological trait can be as great between individuals within a species as between
species. We therefore propose an individual based approach to examine the functional
attributes of arable plants that affect resource acquisition, partitioning and energy
transfer through the food web. Capsella (shepherds purse) is proposed as a key species
that is widespread, ecologically important, and physiologically and genetically diverse.
Current understanding of Capsella systematics is therefore reviewed and a
methodological approach is described that establishes a foundation for studies of
biodiversity and function in arable systems.
C. Hawes, G. S. Begg, G. R. Squire and P. P. M. Iannetta, Ecosystem Management and
Biotechnology, Scottish Crop Research Institute, Invergowrie Dundee DD2 5DA
Scotland, UK (chawes@scri.sari.ac.uk).
521
(a)
0
0
5
1930 seed density m 2 (log N+1)
1
2
3
4
1953 seed density m 2 (logN+1)
(b)
0
0
1
2
3
4
1925 seed density m 2 (log N+1)
germination and flowering characteristics. Spring, autumn and year-round germination types are equally
represented, and a third of species have the potential
to flower throughout the year. These common species
therefore possess life history traits that enable them to
persist despite the high levels of disturbance characteristic of arable systems. All of these common species are
adapted to high soil nutrient conditions and a high
proportion (about 40%) are thought to be increasing in
population density (Ecological flora of the British Isles
database, Table 1). By contrast, those classified as rare
(occurring in 2% of sites or less) are less well adapted to
arable conditions and many of these species populations
are declining (Table 1). A greater proportion are
restricted to autumn-germination, have flowering periods of just one to two months a year, and display
preferences for lower fertility soils. Field management
practices therefore select for particular plant traits and
can have a major impact on the range of plant types
represented in the arable seedbank.
OIKOS 109:3 (2005)
Apiaceae
Asteraceae
Aethusa
cynapium
Petroselinum
segetum
Scandix
pectin-veneris
Torilis
arvensis
Centaurea
cyanus
Chrysanthemum
segetum
Cirsium arvense
Filago apiculata
Boraginaceae
Filago
pyramidata
Matricaria
recutita
Senecio vulgaris
Sonchus
oleraceus
Tripleurospermum
inodorum
Lithospermum
arvense
Myosotis arvensis
Chromosome
number
Ploidy level
DNA content
Mycorrhizal
association
Fertilization
/ /
Flowering period
(months)
Germination
Life history
Number of insect
species supported
Competitive
ability
Soil nutrient
requirement
Frequency
(and dynamics)
Species
Family
Common
name
Table 1. A representative list of 66 arable weed species with ecological, physiological and genetic information collated from Marshall et al. (2003), Cussons and Courtney (1995),
Clapham et al. (1962), Naylor (2002), from unpublished seedbank data held at SCRI, and from http://www.rbgkew.org.uk/cvalues/ and http://www.york.ac.uk/res/ecoflora/cfm/ecofl/
index.cfm/. Frequency of occurrence in the 200 sites sampled for the Farm-Scale Evaluations (Champion et al. 2003)-rare (R): B/1%; frequent (F): 1 /50%; common (C): /50% of sites;
(/) indicates declining populations, (/) increasing population, (s) stable populations from references in the ecoflora database. Soil nutrient preferences / very fertile (VF), fertile (F),
infertile (I), very infertile (VI). Competitive ability is classified by competitive index (CI) or numbers of plants per m 2 to cause 5% yield loss: low (L) CI B/0.05//50 plants per m 2;
medium (M) CI/0.5 /1/10 /50 plants m 2; High (H) CI/1/ B/10 plants m 2. Life history is summarized as A (annual), B (biennial) or P (perennial). Germination is predominantly Sp
(spring), Au (autumn), W (winter) or Y (all year round). Flowering period is given as the number of months a year in which the species has the potential to flower. Fertilization is
described as primarily outcrossing (X) and/or selfing (S). Significance of each species to microbial communities is indicated in terms of their known mycorrhizal associations, where 0/no
known association, 1/rarely, 2/occasionally and 3/normally mycorrhizal. Genetic information is summarised as the 1C nuclear DNA content, ploidy level and chromosome number
of each species. / indicates that no data were available for the species.
fools parsley
/
/
Sp
7 /8
5 /125
1.64
X, S
/
/
20
corn parsley
/
/
Au
8 /9
30 /100
2.2
/
/
/
/
18
shepherds-needle
R (/)
/
/
Au, (Sp)
4 /7
15 /50
/
/
/
16
spreading
hedge-parsley
corn flower
R (/)
/
/
Au
7 /9
/
/
/
/
2.05
12
R (/)
F-I
/
/
Au, Sp
6 /8
20 /90
2.6 /4.5
/
/
24
corn marigold
R (/)
/
/
Au, (Sp)
6 /8
20 /60
1.9
X, S
7.35
18
creeping
thistle
red-tipped
cudweed
broad-leaved
cudweed
scented
mayweed
groundsel
smooth
sow-thistle
scentless
mayweed
field gromwell
VF-F
50
Au
7 /9
30 /90
1.2
X, S
1.55
34
/
/
/
/
7 /8
/
/
/
/
/
/
/
VI
/
/
/
/
/
5 /30
0.05
/
/
/
/
/
C (/)
15
6 /7
15 /60
0.06
/
/
/
/
/
C (s)
C (/)
VF-VI
F
L
/
47
28
A
A
Y
Sp
1 /12
6 /8
8 /45
20 /150
0.22
0.27
S
S
2
3
1.58
1.60
4
4
40
32
C (/)
31
7 /9
15 /60
0.29
/
1.95
/
18
VF-F
/
/
Au, (Sp)
5 /7
30 /80
6.0
/
/
/
28
VF-F
Au, (Sp)
4 /9
15 /60
0.29
/
/
24/48/52
field forgetme-not
523
524
Brassicaceae
Caryphyllaceae
Alyssum
alyssoides
Arabidopsis
thaliana
Capsella
bursa-pastoris/
rubella
Sinapsis arvensis
Cerastium
fontanum
Silene gallica
Silene noctiflora
Spergula arvensis
Stellaria media
Chenopodiaceae
Euphorbiaceae
Fabaceae
Fumariaceae
Chenopodium
album
Euphorbia
helioscopia
Euphorbia
platyphyllos
Vicia tenuissima
Fumaria bastardii
Fumaria
officinalis
Fumaria purpurea
Geraniaceae
Lamiaceae
Geranium
dissectum
Ajuga
chamaepitys
Galeopsis
augustifolia
small Alison
I-VI
/
/
5 /6
7 /25
0.56
X, S
/
/
thale cress
F-VI
/
Au
4 /5
5 /50
0.03
shepherds purse
/
13
1 /12
3 /40
0.1
X, S
charlock
common
mouse-ear
smallflowered catchfly
nightflowering
catchfly
corn spurrey
common
chickweed
fat-hen
F (/)
F (s)
VF-F
F-I
M
/
38
22
A
P/A
Sp
Au
5 /7
4 /9
30 /80
45
2.0
0.16
R (/)
/
/
Sp
6 /10
15 /45
R (/)
VF-F
/
/
Sp
7 /9
15 /45
F (/)
C
F
F
/
M
/
71
A
A
Sp, (Au)
Au, Sp
6 /9
1 /12
C ( /)
31
Sp, (Au)
sun spurge
VF-F
/
/
broad-leaved
spurge
slender tare
tall rampingfumitory
common
fumitory
purple
rampingfumitory
cut-leaved
cranes-bill
ground- pine
VF-F
/
/
R
R (s)
/
VF-F
/
/
VF-VI
red hemp-nettle
Chromosome
number
Ploidy level
DNA content
Mycorrhizal
association
Fertilization
/ /
Flowering period
(months)
Germination
Life history
Number of insect
species supported
Competitive
ability
Soil nutrient
requirement
Frequency
(and dynamics)
Species
Family
Common
name
Table 1 (Continued )
/
32
0.2
2/4
10
0.7
32
X, S
X, S
/
1
0.38 2
2.93 16
18
144
0.5
/
/
/
24
0.92
/
/
/
/
24
5 /60
5 /40
0.42
0.37 /0.67
X, S
X, S
1
2
1.05
1.05
/
7
18
42
7 /10
10 /100
0.77 /1.5
X, S
2.33
54
Sp
5 /10
10 /50
2.49
/
/
42
Au, (Sp)
6 /10
15 /80
/
/
/
/
18/28
/
/
A
A
/
Au, Sp
6 /8
4 /10
/
100
/
/
/
S
/
/
/
/
/
/
14
/
/
/
Sp, (Au)
5 /10
100
/
X, S
/
/
28
F-I
/
/
/
7 /10
100
/
/
/
/
/
VF-F
Au, Sp
5 /8
10 /60
/
/
22
VI
/
/
Au
5 /9
5 /20
1.16
/
/
/
/
28
R (/)
/
/
Sp
7 /10
10 /80
5.4
X, S
/
/
/
16
2.0 /3.0
Galeopsis tetrahit
Papaveraceae
Poaceae
Lamium
purpureum
Papaver
argemone
Papaver hybridum
Papaver rhoeas
Alopecurus
myosuroides
Anisantha sterilis
Apera spica-venti
Avena fatua
Elytrigia repens
Phleum phleoides
Poa annua
Polygonaceae
Primulaceae
Fallopia
convolvulus
Persicaria
maculosa
Polygonum aviculare
Polygonum
rurivagum
Rumex
obtusifolius
Anagallis arvensis
Chromosome
number
Ploidy level
DNA content
Mycorrhizal
association
Fertilization
/ /
Flowering period
(months)
Germination
Life history
Number of insect
species supported
Competitive
ability
Soil nutrient
requirement
Frequency
(and dynamics)
Common
name
Species
Family
Table 1 (Continued )
common hempnettle
red dead-nettle
F (///)
VF-F
/
Sp
7 /9
10 /100
4.83
/
/
32
C (/)
/
Au, Sp
3 /10
10 /45
0.7
X, S
1.10
18
prickly poppy
R (/)
I-VI
/
/
Au, Sp
6 /7
10 /50
0.15
X, S
/
/
/
12/42
rough poppy
common
poppy
black grass
R
F (/)
F-I
F-VF
/
M
/
8
A
A
Au, Sp
Au, (Sp)
6 /7
6 /8
10 /50
10 /50
0.16
0.14
S
X
/
1
2.48
2.63
2
2
14
14
C ( /)
VF-F
Au, (Sp)
5 /9
20 /70
2.0
4.33
/
14
barren brome
loose silky-bent
winter wild oat
Couch grass
purple-stem
cats-tail
annual
meadow grass
black
bindweed
redshank
F (/)
R
F (/)
F (/)
R
VF-F
/
VF-I
F
/
M
/
H
/
/
/
/
4
/
/
A
/
/
P
P
Au
Au
(Au), Sp
Au, Sp
/
5 /7
6 /7
7 /9
6 /9
6 /7
5 /100
20 /100
30 /100
30 /120
10 /40
8.4
0.12
14.1
2.02
/
S
X
S
X
X
/
3
2
3
3
3.35
5.40
14.15
/13.0
/
/
2
6
/
/
14
14
42
42
14/28
C (/)
52
1 /12
5 /30
0.48
2.88
28
/
/
Sp
5 /10
30 /120
1.28
0.72
40
F (/)
/
20
Sp
6 /10
25 /75
2.05
X, S
/
/
knotgrass
cornfield
knotgrass
broad leafed
dock
scarlet
pimpernel
C (/)
R
F
I
L
/
61
/
A
A
Sp
Sp
7 /10
7 /10
1.69
/
S
S
1
/
0.85
/
4
/
60
60
F (/)
VF-F
/
/
Sp
6 /10
15 /120
1.10
1.53
40
Sp
6 /8
6 /30
0.4
/
1.75
40
pheasants eye
corn buttercup
R (/)
R (/)
F-VI
I-VI
L
/
/
/
A
A
Sp, (Au)
Au
7
6 /7
10 /40
15 /60
/
X
/
/
/
6.13
/
4
32
32
common
cleavers
corn cleavers
F (/)
30
Au, (Sp)
6 /8
15 /120
7.25
X, S
1.03
/
44
/
/
/
/
6 /9
/
/
/
/
/
/
44
200
30
Ranunculaceae
Rubiaceae
Adonis annua
Ranunclus
arvensis
Galium aparine
525
Galium
tricornutum
4.8 /18.4
12
526
Scrophulariaceae
Melampyrum arvense
Veronica persica
Veronica praecox
Solanaceae
Urticaceae
Valerianaceae
Violaceae
Veronica
triphyllos
Solanum nigrum
Urtica dioica
Urtica urens
Valerianella
dentate
Valerianella
rimosa
Viola arvensis
field
cow-wheat
common
field-speedwell
brecklandspeedwell
fingered
speedwell
black
nightshade
common nettle
small nettle
narrow-fruited
cornsalad
broad-fruited
cornsalad
field pansy
R (/)
VF-I
/
/
Sp
6 /9
20 /60
Au, Sp
1 /12
10 /40
I-VI
/
/
/
3 /6
R (/)
VI
/
/
/
3 /6
VF-F
/
Sp
7 /9
C (/)
C
R (/)
F
F
F /I
/
/
L
/
/
/
P
A
A
Sp
Sp, (Au)
Au, Sp
/
VF-F
18.7
Chromosome
number
Ploidy level
DNA content
Mycorrhizal
association
Fertilization
/ /
Flowering period
(months)
Germination
Life history
Number of insect
species supported
Competitive
ability
Soil nutrient
requirement
Frequency
(and dynamics)
Species
Family
Common
name
Table 1 (Continued )
X, S
/
/
18
0.52
X, S
0.78
28
5 /20
/
/
/
/
/
18
5 /20
/
X, S
/
/
/
14
60
0.97
3.10
72
6 /8
4 /10
6 /7
30 /150
10 /60
7 /40
0.2
0.5
1.0
X
/
S
2
0
/
1.58
0.33
/
4
/
/
52
24/26/52
14
/
7 /8
7 /40
/
/
/
/
14
Au, Sp
4 /10
15 /45
0.2
/
/
34
5.2
4.6
4.0
3.4
2.8
2.2
19151930
19401944
19531962
19661978
19891999
20002003
albeit at reduced density. Species richness might therefore be recoverable by improved land management, such
as: reducing herbicide use (Cussons and Courtney 1994,
Young et al. 2001), altering the timing and frequency of
herbicide application (Champion et al. 2003), and
shifting the balance in favour of spring-sown crops in
the rotation (Squire et al. 2000, Young et al. 2001).
Many of the species whose populations can be
enhanced have an important role in supporting a diverse
arthropod community. Although data on the insect
associations for arable weeds is only available for 60%
of the species listed in Table 1, many are known to
support a wide range of insect taxa. Species that are
thought to be beneficial to higher trophic levels include
members of the Chenopodiacea, Compositae, Leguminosae, Polygonaceae and Umbelliferae (Altierri 1999,
Wilson et al. 1999, Marshall et al. 2003). An understorey
of these types of species could also have positive impacts
on crop productivity and health through reduction in
pest and disease populations. In contrast, some of the
most competitive weeds that are targets of weed control,
such the competitive grass weeds Alopecurus myosuroides and Bromus sterilis, provide a lower quality of
resource and may therefore be less important to invertebrate consumers (Table 1).
100
10
0.1
0
20
40
60
80
100
120
140
Individuals can be characterized at both the phenotypic and molecular level, since genes have been identified that regulate patterns of resource allocation and lifehistory tradeoffs (Mitchell-Olds 1996). A suite of
methods is available to study these properties in crop
species for the development of new crop varieties.
Accessions are gathered that represent diverse phenotypes from different climatic and geographic regions.
These are cultured in controlled or standard conditions
and characterized using genetic markers capable of
identifying individuals with desirable phenotypic traits.
Transfer of these common agronomic approaches to
arable weeds would allow the genetic basis for observed
phenotypic variation to be quantified. This is necessary
to determine the heritability of expressed traits, thereby
placing measures of functional diversity in an evolutionary context. However, to achieve this, information is
required on the structure of the genome, the mating
system, the genetic distinctness of individuals (as indicated by genetic marking techniques), and the shift in
trait values that occurs through cross-breeding. The
genetic, physiological and ecological attributes of 66
representative arable species, summarised in Table 1, are
used for illustration below.
Conclusions
The important traits for survival of arable plants, given
the high levels of disturbance within fields, are those that
relate to germination, flowering time, seed production
and seedbank persistence. The spread of these traits
among individuals in an arable community will determine the level of functional diversity and the amount of
different resource types that are available for consumers.
The abundance and diversity of arable weeds may be
enhanced with little impact on crop yield if field
management practices can be designed that select for
weeds with particular characteristics: high resource
acquisition and utilisation efficiency, low nutrient and
light requirements, a spread of germination and flowering periods within the window between disturbance
events, and high resource quality for invertebrate and
other consumers. If within-field communities with these
characteristics are to be achieved, detailed information is
required on the ecological, physiological and genetic
traits of arable weeds. However, there are major gaps in
current knowledge for even the most common species.
Even less is known about the degree of variation within
these species that contributes to overall levels of functional diversity. We therefore propose an individual,
trait-based approach to define functional diversity in
arable systems using, in the first instance, Capsella as the
target species. Systematic investigation of phenotypic
and genotypic variation at the individual level is now
required to understand the mechanisms underlying the
maintenance of diversity in productive arable systems.
This can be achieved through phenotypic measurements
of individual plants, exhaustively sampled from the
seedbank at a range geographic locations with varying
management histories, and grown ex situ in a controlled
environment from seed germination to death (Gaudet
and Keddy 1988). These measurements should be
associated with genetic characterisation of individuals
of known phenotypes using molecular markers for key
life history strategies and trade-offs. If successful, it will
be necessary to extend this approach from Capsella to
other species that collectively account for most of the
primary productivity of arable ecosystems. Quantifying
functional diversity according to variation in traits
between individual plants will provide the basis for
future assessment of the impact of biodiversity on
system functions such as resilience and productivity.
OIKOS 109:3 (2005)
References
Aarssen, L. W. and Taylor, D. R. 1992. Fecundity allocation in
herbaceous plants. / Oikos 65: 225 /232.
Aarssen, L. W. and Jordan, C. Y. 2001. Between-species patterns
of covariation in plant size, seed size and fecundity in
monocarpic herbs. / Ecoscience 8: 471 /477.
Acarkan, A., RoBerg, M., Koch, M. et al. 2000. Comparative
genome analysis reveals extensive conservation of genome
organisation for Arabidopsis thaliana and Capsella rubella .
/ Plant J. 23: 55 /62.
Aksoy, A. 1996. Autecology of Capsella bursa-pastoris. / Univ.
of Bradford.
Aksoy, A., Dixon, J. M. and Hale, W. H. G. 1998. Capsella
bursa-pastoris (L.) Medikus. (Thlaspi bursa-pastoris (L.),
Bursa bursa-pastoris (L.) Schull, Bursa pastoris (L.) Weber).
/ J. Ecol. 86: 171 /186.
Aksoy, A., Hale, W. H. G. and Dixon, J. M. 1999. Towards a
simplified taxonomy of Capsella bursa-pastoris (L.) Medik.
(Brassicaceae). / Watsonia 22: 243 /250.
Almquist, E. 1921. Bursa pastoris. / Supplement to the report
of the Botanical Society and Exchange Club of the British
Isles 6: 179 /207.
Almquist, E. 1923. Studien uber Capsella bursa-pastoris II.
/ Acta Hortii Bergiani 7: 41 /95.
Almquist, E. 1929. Zur Artbildung in der reien Natur. / Acta
Horti Bergiani 9: 37 /75.
Altierri, M. A. 1999. The ecological role of biodiversity in
agroecosystems. / Agric. Ecosyst. Environ. 74: 19 /31.
Andow, D. A. 1991. Vegetational diversity and arthropod
population response. / Annu. Rev. Entomol. 36: 561 /586.
Arabidopsis Genome Initiative 2000. Analysis of the genome
sequence of the flowering plant Arabidopsis thaliana .
/ Nature 408: 796 /815.
Asteraki, E. J., Hanks, C. B. and Clements, R. O. 2003. The
influence of different types of grassland field margin on
carabid beetle (Coleoptera, Carabidae) communities.
/ Agric. Ecosyst. Environ. 54: 195 /202.
Barot, S. and Gignoux, J. 2004. Mechanisms promoting plant
coexistence: can all the proposed processes be reconciled?
/ Oikos 106: 185 /192.
Bengtsson, J., Fagerstrom, T. and Rydin, H. 1994. Competition
and coexistence in plant communities. / Trends Ecol. Evol.
9: 246 /250.
Bennett, M. D. and Smith, J. B. 1976. Nuclear DNA amounts in
angiosperms. / Philos. Trans. R. Soc. Lond. 274: 227 /274.
Blazquez, M., Koornneef, M. and Putterill, J. 2001. Flowering
on time: genes that regulate the floral transition. / EMBO 2:
1078 /1082.
Bolnick, D. I., Svanback, R., Fordyce, J. A. et al. 2003. The
ecology of individuals: incidence and implications of individual specialization. / Am. Nat. 161: 1 /28.
Bown, J. L. 2000. Issues of scale in individual-based models:
applications in fungal and plant community dynamics.
/ PhD thesis, Univ. of Abertay, Dundee, UK.
Brenchley, W. E. 1918. Buried weed seeds. / J. Agric. Sci. 9: 1 /
31.
Brenchley, W. E. 1920. Weeds of farmland. / Longmans, Green
and Co.
Brenchley, W. E. and Warington, K. 1933. The weed seed
population of arable soil. II Influence of crop, soil and
methods of cultivation upon the relative abundance of viable
seeds. / J. Ecol. 21: 103 /127.
Buckelew, L. D., Pedigo, L. P., Mero, H. M. et al. 2000. Effects
of weed management systems on canopy insects in herbicide
resistant soybeans. / J. Econ. Entomol. 93: 1437 /1443.
531
532
533
534