See

discussions, stats, and author profiles for this publication at:

https://www.researchgate.net/publication/229542529

Macrophytes: Ecology of Aquatic Plants

Chapter · September 2009

DOI: 10.1002/9780470015902.a0020475

CITATIONS READS

9 271

2 authors:

Gudrun Bornette Sara Puijalon

French National Centre for Scientific Re… Claude Bernard University Lyon 1
96 PUBLICATIONS 3,255 CITATIONS 16 PUBLICATIONS 607 CITATIONS

SEE PROFILE SEE PROFILE

All in-text references underlined in blue are linked to publications on ResearchGate, Available from: Gudrun Bornette
letting you access and read them immediately. Retrieved on: 11 April 2016
Macrophytes: Ecology of Advanced article

Aquatic Plants . Introduction

Article Contents

Gudrun Bornette, CNRS, Ecology of Fluvial Ecosystems Université Lyon1, Villeurbanne Cedex, . Classification of Macrophytes

. Challenges and Adaptations to Life in Water

France
. Reproduction and Dispersal of Aquatic Plants
Sara Puijalon, Netherlands Institute of Ecology (NIOO-KNAW), Centre for Estuarine and Marine . Abiotic Parameters that Govern the Occurrence of
Macrophytes
Ecology, Yerseke, The Netherlands
. Biotic Interactions
. Consequences for the Structure and Dynamics of Plant
Aquatic plants contribute to maintaining key functions and related biodiversity in Communities
freshwater ecosystems, and to provide the needs of human societies. The way the . Threats to Macrophytes
ecological niches of macrophytes are determined by abiotic filters and biotic ones is Online posting date: 15th September 2009
considered. A simple, broadly applicable model of the distribution of growth forms
according to abiotic filters is proposed. The consequence for the dynamics of plant
communities and the main threats to macrophyte occurrence and diversity are
discussed.

Introduction Classification of Macrophytes

Macrophytes are primary producers and provide habitats
and refuges for periphyton, zooplankton, other inverte-
brate species, fish and frogs. They also play key functions in
biogeochemical cycles (e.g. organic carbon production and
phosphorous mobilization) directly influencing hydrology
and sediment dynamic of freshwater ecosystems through
their effects on water flow. Macrophytes contribute to
maintain key functions and related biodiversity in fresh-
water ecosystems, but also through the services they
provide directly to man. See also: Wetlands
The aim here is to present the main processes that
determine macrophyte distribution, community composi-
tion and biodiversity, and plant succession in temperate
freshwaters. Only submerged macrophytes that live
permanently or most of the time under water are
considered, as such species have rather contrasting habitats
and specific requirements compared to helophytes (i.e.
partly emerged macrophytes). In terms of habitats, the
focus will be on natural habitats of freshwater macro-
phytes, that is, continental standing and running ecosys-
tems, as lakes, ponds, wetlands, rivers and streams. Species
of brackish waters are not considered, as the requirements
and stress that affect seagrasses partly differ from those
affecting freshwater species.
Macrophytes include bryophytes (mosses, liverworts and
hornworts), pteridophytes (ferns and fern allies) and
spermatophytes (seed-bearing plants) whose photosynthe-
tically active parts are permanently submerged in water or
which float on the surface of water (Cook, 1999). Less than
2% of vascular plants are aquatic. There are similar
proportions of aquatics in monocots and dicots at the level
of the superorder, but proportionally more aquatics in the
monocots than in the dicots at the family level (35% of
monocot families present aquatic species, but only 12.1% of
dicot families). Only a few groups of Angiosperms are fully
aquatic, as for example, Nympheales, Hydrocharitales,
Zosterales, Alismatales and Podostemales. Most other
aquatic species belong to groups where only some families
or some genera are aquatic. The question of the aquatic
character of the first angiosperms has been heavily debated,
but so far has not been elucidated. The recent angiosperm
classification outlines the basal lineage of several aquatic
taxa (e.g. Ceratophyllales, Nympheaceae and Alismatales;
Angiosperm Phylogeny Group, 2003), and fossil records
suggest the early colonization of aquatic habitats by
angiosperms. In addition, angiosperms were able to return
several times to water in the course of evolution.

Challenges and Adaptations to Life in

Water
ELS subject area: Ecology
An angiosperm has to solve several problems to successfully
How to cite: colonize aquatic habitats. The low-diffusion rate of gases
Bornette, Gudrun; and, Puijalon, Sara (September 2009) Macrophytes: (and more particularly carbon dioxide and oxygen) leads to
Ecology of Aquatic Plants. In: Encyclopedia of Life Sciences (ELS). John
problems of carbon and oxygen supply. The mobilization
Wiley & Sons, Ltd: Chichester.
DOI: 10.1002/9780470015902.a0020475
and transport of nutrients in the plant are further constraints,
as transpiration from leaves can no longer drive internal

ENCYCLOPEDIA OF LIFE SCIENCES & 2009, John Wiley & Sons, Ltd. www.els.net 1
 Macrophytes: Ecology of Aquatic Plants
water transport in aquatic environments, except for plants
with floating leaves. Among factors that can act positively on
plant growth, the high density of water (more than 700 times
higher) compared to air allows plants to be mechanically
supported. Nutrients occur in water, in contact with the
aboveground organs of the plants and are potentially
available for plant growth. Aquatic plants present a common
set of traits that adapt them to such habitats (Rascio, 2002).
Among them, the occurrence of aerenchyma increases
oxygen flux from shoots to roots. A greater leaf surface
(when the same species is able to produce both emerged and
submerged leaves, the leaf surface is strongly greater for
submerged leaves compared to emerged ones), together with
a thin cuticle, increase contacts with water and carbon
uptake. Water plants are usually poorly lignified, as water
protects plants from gravitational stress, and xylem cells are
replaced by a few xylem lacunae, with a much higher
hydraulic conductance. The occurrence of a casparian band
(band of cell wall material composed of lignin and/or
suberin, in the radial and transverse walls of the endodermis)
in the endodermis of the stalk and the leaves is in this context
viewed as a hydraulic insulator, preserving the passive
diffusion of nutrient solutions towards the aerenchyma, and
suggesting that root pressure is the major driving force for
water and nutrients from roots to apical meristems. Poor
oxygen availability underwater accounted for the evolution
of large aerenchyma throughout the plant to improve the
photosynthetic oxygen flux from shoots to roots buried in
anoxic sediments and to the neighbouring rhizosphere.
Reproduction and Dispersal of Aquatic
Plants
Many aquatic species are phylogenetically descended
from terrestrial plants that later adapted to aquatic life.
As a consequence, many species still have aerial se-
xual reproduction, which increases the risk of reproduction
failure through submersion by water movements. As
a consequence, insect-pollinated species are also frequently
able to propagate by vegetative means (rhizome ex-
pansion, or stem fragmentation). For example, Batrachium
sp. (Ranunculaceae) or Hottonia sp. (Primulaceae) have
aerial flowers, but have shown a very high capacity for
vegetative growth. In some groups, the flower petals have
regressed, suggesting a tendency towards wind pollination
(e.g. Potamogetonaceae), increasing the reproduction
success when there is a lack of insects. Some taxa have
developed submerged reproduction (e.g. Najadaceae)
achieved by water pollination (e.g. through filiformis
pollen grains or grains with reduced exine structure,
dispersed by water; Ackerman, 2000; Tanaka et al., 2004)
and frequently produce large amount of seeds. A drop in
water level usually favours sexual reproduction, and the
composition of plant communities in temporary ecosys-
tems relies heavily on the ability of species to produce
viable seeds.
2 ENCYCLOPEDIA OF LIFE SCIENCES & 2009, John Wiley & Sons, Ltd. www.els.net
Most aquatic species are able to propagate and multiply
vegetatively. However, the ability of regrowth from plant
fragments differs among taxonomic groups (Barrat-Segre-
tain et al., 1998). For example, evergreen species are usually
more efficient at propagating by vegetative means, because
they are able to grow during the winter in cool water
situations (e.g. Luronium natans and Berula erecta).
Floods are a major means of dispersal for aquatic plants.
The efficiency of dispersal depends on seed buoyancy and
on the ability of plants to break themselves up and re-
grow from broken dispersed fragments. Dispersal effi-
ciency depends on river discharge (Boedeltje et al., 2004),
probably because river discharge is related to flow velocity
(and thus to the magnitude of the hydrodynamic forces and
the plant capacity to resist to breakage and uprooting) and
to the surface inundated by water (and thus on the
efficiency of water for capturing seeds that are released
along the banks). However, vegetative fragments are 10
times more numerous than seeds in drifting material,
emphasizing the key role of vegetative growth in short and
long-distance dispersal of aquatic plants. Some species
have been shown, at least in some geographical areas,
to disperse almost exclusively by vegetative means (e.g.
Utricularia australis f. australis; Araki and Kadono, 2003).
Many shorebirds and water birds are important consumers
of seeds of aquatic plants and also contribute to water
plant dispersal (Green et al., 2002), essential in the case of
low-hydraulic connectivity among water bodies.
Abiotic Parameters that Govern the
Occurrence of Macrophytes
Parameters related to plant nutrition
Water transparency
The occurrence of macrophytes is strongly governed by
water transparency, itself determined by the colour of the
water (partly related to dissolved organic matter), the
concentration of suspended solids and the concentration of
plankton. The depth limit of macrophytes is when the water
transparency allows less than 1–4% light to reach the
plants (Sculthorpe, 1967). Angiosperms are confined to
shallower depths than bryophytes and charophytes, and
the minimum light requirement of submersed macrophytes
is linked to their growth form, as this determines plant
biomass per unit of light-absorbing surface area, that is, the
ability of plants to harvest light (Middelboe and Markager,
1997). Because they are frequently characterized by a low-
growth rate, rosette growth forms are limited to clear water
lakes or shallow zones (Vestergaard and Sand-Jensen,
2000), whereas caulescent angiosperms, able to produce
erected stems that grow towards the water surface are able
to colonize deeper waters (Figure 1). In turbid situations,
plant communities are dominated by species with floating
leaves, associated with free-floating species if the nutrient
content of the water is sufficient to allow plant growth.
 Macrophytes: Ecology of Aquatic Plants

Small Small Tall Tall species with Free-floating

rosettes caulescents caulescents floating leaves species

Growth
forms

Examples of Samolus sp. Callitriche sp. Myriophyllum sp. Nymphea sp. Lemna sp.
taxonomic units Berula sp. Elodea sp. Potamogeton Potamogeton Hydrocharis sp.
lucens nodosus
Water
+
transparency −

Seasonal
fluctuations in − +
temperatures
Availability of
carbon dioxide +
and oxygen −

Substrate Coarse or
grain-size cohesive − Indifferent

Nutrient
level − +

Water +
movements −

Figure 1 Expected distribution of the five major growth forms of aquatic plants along several gradients of abiotic factors. For example, for water transparency,
species with small rosettes tend to occur preferably in sites with high transparency, whereas free-floating plants will occur more frequently in water with low
transparency.

Water temperature
Freshwater ecosystems are usually considered to be more
stable in terms of temperature variations than terrestrial
ecosystems. Most aquatic plant species exhibit optimal
rates of photosynthesis at relatively high temperature
(between 208C and 358C and between 28 and 328C,
depending on the author). However, some species are able
to grow below 108C (Boylen and Sheldon, 1976). Several
aquatic species are evergreen or wintergreen and poten-
tially grow in ecosystems with reduced seasonal fluctua-
tions, such as groundwater-supplied ecosystems, due to the
maintenance of relatively high winter temperatures, but
present relatively prostrate growth forms in winter (e.g.
rosette or small caulescents). See also: Stream and River
Community Structure
Water nutrient content
Carbon, nitrogen and phosphorous are the main nutrients
required for plant life, and their availability therefore
governs primary production in freshwater ecosystems.
Two groups of species can be distinguished, those that can
only use carbon dioxide (in air, water or in sediment) and
those that can use both carbon dioxide and HCO32. Plants
with floating leaves are able to use carbon dioxide from the
air. Among submerged species, some are restricted to
carbon dioxide (Maberly and Madsen, 1998). Isoetids, that
is, small evergreen plants with inherently slow growth, use
sediment carbon dioxide (produced by respiration of roots
and animals), and occur in nutrient-poor, freshwater
habitats, have a low photosynthetic rate and poor capacity
for carbon extraction compared to other macrophyte
species. Submerged species able to use HCO2 3 tend to grow
in alkaline waters and to have a higher relative growth rate.
Several studies demonstrated that phosphorous and
ammonium concentrations in water govern the composi-
tion of aquatic plant communities, and indices link the
composition of macrophyte communities and nutrient
levels in the water (Schneider, 2007). On the one hand, free-
floating macrophytes absorb their nutrients from water,
and are thus usually found in ecosystems where the nutrient
level in the water column is high. Rooted plants, on the
other hand, can use nutrients both in the water column and
in the substrate. Tall species tend to occur in nutrient-rich
waters. Small species (such as species with small rosettes
and caulescent species) colonize water with lower nutrient
levels, the smallest species being restricted to the clearest
nutrient-poor waters (Figure 1). Nutrient levels also affect

ENCYCLOPEDIA OF LIFE SCIENCES & 2009, John Wiley & Sons, Ltd. www.els.net 3
 Macrophytes: Ecology of Aquatic Plants
species richness, the highest species diversity observed for
intermediate nutrient levels. Under very low or very high
nutrient levels, either only stress tolerators (low-nutrient
levels) or very competitive species (high-nutrient levels)
dominate. See also: Competition; Eutrophication of Lakes
and Rivers
Substrate characteristics
A few species that grow in running waters are strictly linked
to rocky substrates, and some others are able to anchor
themselves firmly in gravels, and tolerate high-flow velocity
(e.g. approximately 0.5 cm s21). However, most species
require finer substrates. Some grow preferentially on
mineral substrates, usually found along exposed river or
lake banks. Such mineral substrates tend to be highly
cohesive, favouring the anchorage of shallowly rooted
species such as those with small rosettes (Greulich et al.,
2000), but are potentially less favourable for the growth
of tall plants, because they are frequently nutrient-poor.
Tall species tend to grow in fine, usually nutrient-rich
substrates, and are often associated with less stressful
situations, because sediment movements cause plant
uprooting in the most exposed situations (Schutten et al.,
2005). Free-floating species are generally indifferent to
substrate grain size (Figure 1). Substrate cohesive strength
also affects species recruitment. Indeed, high-density seeds
tend to sink in low-density substrates, and cannot be
recruited. Reciprocally, the ability of some species to root
can be significantly reduced in highly cohesive sediment,
restricting their establishment to habitats with low cohesive
substrates (e.g. Najas marina). The dewatering of sediments
with low cohesive strength can increase their density
(Redding and Devito, 2006). Such density increases
contribute to altering very significantly the recruitment of
aquatic species after the dry period, potentially increasing
plant recruitment and growth. Rooted aquatic plants are
also able to take up dissolved nutrients (e.g. phosphorous
and nitrogen) from sediments (Barko and Smart, 1986). In
submerged substrates, the mineralization of organic matter
requires oxygen, and this process leads to a greater demand
for oxygen within the soil and hence stress on aquatic plant
roots. This oxygen depletion in the rhizosphere induces a
series of soil chemical changes, such that the accumulation
of several compounds that can be phytotoxic (Pezeshki,
2001). Such phenomenon can have a major effect on the
survival and recruitment of aquatic species, and select
species that either use atmospheric oxygen as an alternative
source of oxygen, or have adaptations for avoiding radial
oxygen losses (Visser et al., 2000) or for improving oxygen
concentrations and subsequent microbial activity in the
rhizosphere. See also: Rhizosphere
Water movements
The main direct effects of water movements are the
hydrodynamic force it exerts on rooted macrophytes.
Those forces scale with plant size and the magnitude of
the different forces depends on the nature of the water
4 ENCYCLOPEDIA OF LIFE SCIENCES & 2009, John Wiley & Sons, Ltd. www.els.net
movements (direction and temporal variability). The
ability for rapid reconfiguration (i.e. changes of plant
shape, alignment of shoots and compaction of leaves) and
high breaking and anchorage strength are the main
adaptations to resisting mechanical damages (Schutten
et al., 2005). In rivers and streams, streamlined shapes can
be very efficient in reducing plant drag, in contrast to wave-
swept environments where plants experience flows from
different directions. In both situations, rosette growth
forms with strap-like leaves or caulescent growth forms
with flexible stems and small leaves are efficient adapta-
tions. Species tend to present small or even dwarfed growth
forms that partly escape flow stress, and greater allocations
to belowground organs and to strengthening tissues,
increasing resistance to breakage and uprooting (Figure
1). Exposure to currents alters and sometimes impedes
plant sexual reproduction. See also: Stream and River
Community Structure
Water movements exert indirect effects on macrophytes.
Particle and nutrient resuspension induced by water
movements potentially limit light penetration in the water
and subsequent plant growth and recruitment. Under
stagnant conditions, photosynthesis can be limited by
diffusion of gas and nutrients through the boundary layer
that develops at the plant surface. A moderate water
stirring reduces the thickness of the boundary layer and
leads to enhanced nutrient and gas fluxes, and hence
improved photosynthesis and plant growth (Crossley et al.,
2002). See also: Stream and River Community Structure
Disturbances
Floods
Depending on the slope and grain size of the riverbed,
floods may either uproot plants and disrupt aquatic plant
communities (and in some cases remove completely fine
sediment deposits), or lead to the deposition of fine
sediment and long-term inundation of wetlands (Bornette
et al., 2008). High frequency and/or intensity of scouring
floods lead to the breakage or uprooting of aquatic plants,
and also wash out fine sediment and seed banks. The
recovery of aquatic plant communities relies on the
occurrence of refuges along the banks of the scoured sites,
and on dispersal of seeds and plant fragments transported
by the flood itself (Henry et al., 1996). In terms of plant life-
history traits, small-sized to intermediate-sized, flexible
growth forms are required, with a high allocation of
resources to anchorage (Figure 2). Long-term flooding has a
rather different impact, such as the long-term submergence
(i.e. usually several weeks) of aquatic plants under turbid
water and sediment deposits that tend to impede plant
growth. Biomass losses through breakage are low, but
plants must resist low light, anoxia and burial (Blom et al.,
1994). The tallest plastic plants with large storage organs
tolerate better long-term submersion the best, as they
can either regenerate vegetative parts of the individ-
uals destroyed by long-term submersion, or elongate
 Macrophytes: Ecology of Aquatic Plants

+ Small + Small
rosettes caulescents

− + − +
Nutrient level

+ + +

Tall species with Free-floating

Tall caulescents
floating leaves species

− + − + − +

Disturbance frequency and/or intensity

Figure 2 Expected distributions of the five major growth forms of aquatic plants according to nutrient levels (mainly phosphates and ammonium in the water)
and flood disturbances. The dark zones of each diagram correspond to the zones that should be more favourable for the group considered. For example, species
with small rosettes tend to occur preferably in sites with low to intermediate nutrient levels but can colonize contrasting sites for disturbance levels.

themselves to reach the light. However, hydrophytes with
both low reserves and growth rates are usually counter-
selected by such events. Rooted submersed plants are
disfavoured, due to the high turbidity of the water and
phytoplankton growth under eutrophic situations. Species
able to anchor themselves in sediment of low cohesive
strength (Handley and Davy, 2002), or those able to
produce adventitious roots or to spread laterally close to
the surface of the newly deposited sediment or at the
surface of the water should also be favoured (Bornette
et al., 2008). See also: Primary Succession
Drawdowns
Rare drawdowns have contrasting effects on plant com-
munities, depending on the intrinsic ability of species to
tolerate dewatering. In many situations, however, it has
been shown that such rare drawdowns favour ruderal
species with abundant seed banks (Brock et al., 2003).
Some species are able to tolerate the drought event, with
plastic modifications of the plant growth form (e.g. leaf
plasticity, size decrease and tissue modifications; Goliber
and Feldman, 1989). For others, the aboveground biomass
dies, and the species survive through subterranean organs.
Some also flower during the water drop and die, constitut-
ing a dormant seed bank that sprouts when the water level
rises again (Casanova and Brock, 2000). In temporary
wetlands (annually dewatered), the maintenance of aquatic
plants relies mainly on their ability to produce seeds able to
tolerate drought. In wetlands characterized by alternating
dry and wet periods, a cyclical seasonal succession occurs,
characterized by the replacement of submerged species,
by floating-leaf species, and finally emergent ones. Some
variations around this pattern can occur over the years
depending on the hydrological characteristics of the
drawdown event (Van der Valk, 2005). In such situations,
slow-growing species, whose maintenance relies mainly on
vegetative multiplication, are disfavoured.
Biotic Interactions
Competition for light
Competition processes occur both within macrophyte
communities and between macrophytes and microphytes.
In terrestrial habitats, the main parameter that determines
competitive ability for light is plant size, which is strongly
correlated with plant biomass (Tilman, 1982). In aquatic
habitats, the ability to reach light is not directly related to
plant size, but to their ability to produce floating or

ENCYCLOPEDIA OF LIFE SCIENCES & 2009, John Wiley & Sons, Ltd. www.els.net 5
 Macrophytes: Ecology of Aquatic Plants
emerged leaves. In addition, the size of plants is not simply
related to their biomass, since submerged organs (e.g.
leaves) present lower construction costs than floating ones
(Titus and Sullivan, 2001). As a consequence, species that
present very contrasting growth forms are considered as
competitively dominant: free-floating species and tall
species with floating leaves (Bornette et al., 1994). Among
other groups, species able to produce through plastic
modifications, either emerged or floating leaves (e.g. many
Alismataceae), are more able to tolerate competition
pressure. Evergreen and wintergreen species (rosette and
small caulescent species) are also able to escape competi-
tion from summergreens (e.g. tall species with floating
leaves and free-floating species) by growing during winter
in ecosystems where there are reduced seasonal variations
of temperature (Greulich and Bornette, 2003).
Allelopathy
Allelopathy, that is, the production and release of chemical
substances that inhibit the growth of other macrophytes or
microphytes, has been demonstrated to occur among
macrophytes (Gross, 2003), even though many published
studies fail to demonstrate in an unfalsifiable way the
allelopathic mechanism, and the toxic compound involved
in the interaction (Ervin and Wetzel, 2003). These authors
provide a review of the relevant papers that identify
allelopathic species, and show that allelopathic effects of
macrophytes occur in contrasting taxonomic groups such
as Najadaceae or Potamogetonaceae. In some situations,
such mechanisms impede microphyte growth and maintain
a clear water stage in lakes (Hilt and Gross, 2008).
Herbivory
Aquatic plants are consumed by many animals, such as
fishes, birds, mammals, molluscs and arthropods (insects,
crustaceae), and plant populations can also be significantly
affected by grazing (Gross et al., 2001). Some grazers
specialize on one macrophyte species (e.g. the coleoptereae
Galerucella nymphaeae that eat Nuphar leaves), but many
of them are polyphaeous. The sensibility of aquatic plants
to graze depends on taxonomic groups (Elger et al., 2004),
but also on the developmental stage, the submerged vs.
floating type of leaves (Sand-Jensen and Jacobsen, 2002),
and on the organ considered, some species grazing
specifically flowers, and seedlings being more susceptible
to graze than fully developed plants. The occurrence of a
cuticle, or strengthening tissues, decreases the sensitivity of
aquatic plants to graze (Elger and Willby, 2003). Aquatic
plants also produce many chemical compounds (tannins,
phenols, alkaloids), and it has been suggested that some of
them could be allelopathic (Smolders et al., 2003).
Other biotic interactions
Aquatic plants can be colonized by mycorrhizal fungi
(Sraj-Krzic et al., 2006). The colonization levels of some
species able to grow indifferently submerged or as a
6 ENCYCLOPEDIA OF LIFE SCIENCES & 2009, John Wiley & Sons, Ltd. www.els.net
riparian species have been demonstrated to be higher in
terrestrial situations, and highly correlated with plant
phosphorous concentrations, suggesting a positive role of
such fungi for plant phosphorous uptake that could be
increased in dry environments, even though this positive
effect is not significant for all species studied.
Consequences for the Structure and
Dynamics of Plant Communities
Nutrient levels
Nutrient levels govern the occurrence, composition and
diversity of aquatic plant communities. When phosphor-
ous concentration increases, floating species (and among
them several invasive species, such as Echhornia crassipes;
Haller and Sutton, 1973) or microphytes outcompete
submerged species. However, for the same phosphorous
concentration, shallow eutrophic lakes can be dominated
either by phytoplankton, and have a high turbidity, or by
macrophytes, and enjoy relatively clear waters (Scheffer
et al., 1993). In the second case, plant uprooting by
benthivorous fish or zooplankton consumption by zoo-
planktivorous fish are both involved directly (plant
uprooting) and indirectly (decreasing predation on phyto-
plankton) in macrophyte collapse (Jeppesen et al., 1999).
See also: Trophic Cascades
Disturbances
Disturbances also reduced biotic interactions through the
recurrent destruction of part of the communities, decreas-
ing the dominance of the tallest, most competitive species,
favouring ruderal species (e.g. isoetids or small rosettes and
caulescent species; Willby et al., 2000). Plant recolonization
after disturbances is probably more rapid when nutrient
levels are high (Khan and Ansari, 2005), and speeding
up the succession process. As a consequence, the less
competitive species are excluded from the community for a
shorter return time after disturbances in eutrophic situa-
tions than in mesotrophic ones (Figure 2).
Successions
Rates and patterns of change in aquatic plant communities
over time are governed either by the autogenic or allogenic
control of successions. Autogenic successions (i.e. succes-
sions driven by stimuli internal to the ecosystem, in
contrast to allogenic successions, driven by external
stimuli) of water bodies have been described, but many of
these studies are based on synchronic analyses of vegeta-
tion patterns (e.g. along elevation gradients; Appelgren
and Mattila, 2005) and exclude the environmental changes
that potentially modify the succession pattern over time.
Furthermore, they frequently focus on the aquatic–
terrestrial transition, and not on species replacement in
the aquatic phase. As a consequence, the succession of

aquatic macrophytes is rather badly known. A good
example of the first steps of succession is given by the
floristic changes observed in gravel pits and created
wetlands. In such systems, the first colonizers are mainly
seed-dispersed macrophytes (Noon, 1996). Colonization is
also affected by plants that reduce water movements and in
this way stabilize the substrate and reduce turbidity,
potentially favouring further plant recruitment.
Divergent succession patterns may occur depending on
nutrient inputs, and on the environmental changes that
affect water bodies. If nutrient inputs are high, species
richness may decrease, leading to the ultimate disappearance
of macrophytes. If this nutrient enrichment is low or leads to
limited eutrophication (e.g. in calcareous situations with
high groundwater inputs, or in periodically dewatered
wetlands, leading to limited phosphate mobilization;
Lucassen et al., 2005; Smolders et al., 2006), tall competitive
species (such as broad-leafed Potamogeton spp. or Nym-
pheaceae) may progressively dominate plant communities
(Ray et al., 2001). During succession, organic matter
progressively fills in water bodies. During this process, the
habitat characteristics are relatively stable, and few changes
in macrophyte communities occur (Schaminee et al., 2002).
However, the increasing layer of poorly cohesive organic
matter can lead to anchorage difficulties for rooted plants,
and a decrease in plant cover. The first dewatering events
lead to the replacement of strictly aquatic plants by emergent
species or aquatic species with a short life cycle. The
succession would lead more rapidly to the dominance of tall
competitive species (both rooted and floating) under
eutrophic conditions than under oligotrophic ones.
Allogenic processes, as disturbances and other environ-
mental parameters (such as climate fluctuations, ground-
water abstraction) can modify this pattern. In lakes, wave
action and grazing can lead to a shifting mosaic of plant
species, and few changes in species composition can occur
over long periods of time. In smaller water bodies (such as
oxbow lakes or small ponds), disturbances (e.g. dewatering
and floods) also contribute to maintain small poorly
competitive species a presence over long periods of time
(Arts, 2002). Very frequent and intense disturbances can
impede completely the growth of aquatic plants. Inter-
mediately frequent disturbances allow for the maintenance
of a highly diverse shifting mosaic of plant species
(Bornette et al., 1994). In both situations, succession is
slowed down and even stopped (Lippert and Jameson,
1964; van Geest et al., 2005). Low frequency and/or
intensity of flood scouring are unable to impede succession
completely and the subsequent substrate accumulation
leads to the terrestrialization of aquatic ecosystems.
See also: Primary Succession; Secondary Succession
Threats to Macrophytes
The first and frequently described threat is the general
eutrophication of freshwaters, leading to the decrease, and
even the disappearance of submerged species, to the benefit
ENCYCLOPEDIA OF LIFE SCIENCES & 2009, John Wiley & Sons, Ltd. www.els.net
Macrophytes: Ecology of Aquatic Plants
of free-floating species, and ultimately microphytes. The
decrease of natural disturbance regimes in river flood-
plains, due to channelization, dam construction and river
incision, also lead to the disappearance of flood-disturbed
habitats, and associated ruderal species. Groundwater
abstraction for agriculture and human needs also leads to
the disappearance of groundwater-fed aquatic habitats,
and their associated evergreen, poorly competitive species.
Climate change will also lead to increases in average
temperature and water-level fluctuations in most temperate
rivers and lakes. Temperature increases would stimulate
microbial processes, increasing anoxia, ammonium accu-
mulation and related stresses on aquatic plants. Submerged
plants would consequently decrease to the benefit of
potentially invasive species with floating leaves and free-
floating ones. An increase in water-table fluctuations
should favour short-lived species investing strongly in
sexual reproduction, whereas a decrease in such fluctua-
tions should favour strictly aquatic tall clonal species.
Consequently, climate change would lead to the significant
decrease, and even disappearance of oligotrophic and
ruderal species, and should favour the increased abun-
dance of species with floating leaves and free-floating ones,
leading ultimately to a decrease in species richness and
diversity of growth forms in aquatic habitats.
References
Ackerman JD (2000) Abiotic pollen and pollination: ecological,
functional, and evolutionary perspectives. Plant Systematics
and Evolution 222: 167–185.
Angiosperm Phylogeny Group (2003) Classification for the orders
and families of flowering plants. Botanical Journal of the
Linnean Society 141: 339–436.
Appelgren K and Mattila J (2005) Variation in vegetation
communities in shallow bays of the northern Baltic Sea. Aquatic
Botany 83: 1–13.
Araki S and Kadono Y (2003) Restricted seed contribution and
clonal dominance in a free-floating aquatic plant Utricularia
australis R. Br. in southwestern Japan. Ecological Research 18:
599–609.
Arts GHP (2002) Deterioration of atlantic soft water macrophyte
communities by acidification, eutrophication and alkalinisa-
tion. Aquatic Botany 73: 373–393.
Barko JW and Smart RM (1986) Sediment-related mechanisms of
growth limitation in submersed macrophytes. Ecology 67:
1328–1340.
Barrat-Segretain MH, Bornette G and Hering-Vilas-Boas A
(1998) Comparative abilities of vegetative regeneration among
aquatic plants growing in disturbed habitats. Aquatic Botany
60: 201–211.
Blom CWPM, Voesenek LACJ, Banga M et al. (1994) Physiolo-
gical ecology of riverside species: adaptive responses of plants to
submergence. Annals of Botany 74: 253–263.
Boedeltje G, Bakker JP, Ten Brinke A, van Groenendael JM and
Soesbergen M (2004) Dispersal phenology of hydrochorous
plants in relation to discharge, seed release time and buoyancy
of seeds: the flood pulse concept supported. Journal of Ecology
92: 786–796.
7
 Macrophytes: Ecology of Aquatic Plants
Bornette G, Amoros C and Chessel D (1994) Effect of allogenic
processes on successional rates in former river channels. Journal
of Vegetation Science 5: 237–246.
Bornette G, Tabacchi E, Hupp CR, Puijalon S and Rostan J-C
(2008) A model of plant strategies in fluvial hydrosystems.
Freshwater Biology 53: 1692–1705.
Boylen CW and Sheldon RB (1976) Submergent macrophytes:
growth under winter ice cover. Science 194: 841–842.
Brock MA, Nielsen DL, Shiel RJ, Green JD and Langley JD
(2003) Drought and aquatic community resilience: the role of
eggs and seeds in sediments of temporary wetlands. Freshwater
Biology 48: 1207–1218.
Casanova MT and Brock MA (2000) How do depth, duration and
frequency of flooding influence the establishment of wetland
plant communities? Plant Ecology 147: 237–250.
Cook CDK (1999) The number and kinds of embryo-bearing
plants which have become aquatic. Perspectives in Plant
Ecology, Evolution and Systematics 2: 79–102.
Crossley MN, Dennison WC, Williams RR and Wearing AH
(2002) The interaction of water flow and nutrients on aquatic
plant growth. Hydrobiologia 489: 63–70.
Elger A, Bornette G, Barrat-Segretain MH and Amoros C (2004)
Disturbances as a structuring factor of plant palatability in
aquatic communities. Ecology 85: 304–311.
Elger A and Willby NJ (2003) Leaf dry matter content as an
integrative expression of plant palatability: the case of fresh-
water macrophytes. Functional Ecology 17: 58–65.
Ervin GN and Wetzel RG (2003) An ecological perspective of
allelochemical interference in land-water interface commu-
nities. Plant and Soil 256: 13–28.
van Geest GJ, Coops H, Roijackers RMM, Buijse AD and
Scheffer M (2005) Succession of aquatic vegetation driven by
reduced water-level fluctuations in floodplain lakes. Journal of
Applied Ecology 42: 251–260.
Goliber TE and Feldman LJ (1989) Osmotic stress, endo-
genous abcisic acid and the control of leaf morphology
in Hippuris vulgaris. Plant, Cell and Environment 12: 163–
171.
Green AJ, Figuerola J and Sanchez MI (2002) Implications of
waterbird ecology for the dispersal of aquatic organisms. Acta
Oecologica 23: 177–189.
Greulich S and Bornette G (2003) Being evergreen in an aquatic
habitat with attenuated seasonal contrasts – a major compe-
titive advantage? Plant Ecology 167: 9–18.
Greulich S, Bornette G, Amoros C and Roelofs JGM (2000)
Investigation on the fundamental niche of a rare species: an
experiment on establishment of Luronium natans. Aquatic
Botany 66: 209–224.
Gross E (2003) Allelopathy of aquatic autotrophs. Critical
Reviews in Plant Sciences 22: 313–339.
Gross EM, Johnson RL and Hairston NG Jr (2001) Experimental
evidence for changes in submersed macrophyte species compo-
sition caused by the herbivore Acentria ephemerella (Lepidop-
tera). Oecologia 127: 105–114.
Haller WT and Sutton DL (1973) Effect of pH and high
phosphorous concentration on growth of waterhyacinth.
Journal of Aquatic Plant Management 11: 59–61.
Handley RJ and Davy AJ (2002) Seedling root establishment may
limit Najas marina L. to sediments of low cohesive strength.
Aquatic Botany 73: 129–136.
8 ENCYCLOPEDIA OF LIFE SCIENCES & 2009, John Wiley & Sons, Ltd. www.els.net
Henry CP, Amoros C and Bornette G (1996) Species traits and
recolonization processes after flood disturbances in riverine
macrophytes. Vegetatio 122: 13–27.
Hilt S and Gross EM (2008) Can allelopathically active
submerged macrophytes stabilise clear-water states in shallow
lakes? Basic and Applied Ecology 9: 422–432.
Jeppesen E, Jensen JP, Søndergaard M and Lauridsen TL (1999)
Trophic dynamics in turbid and clearwater lakes with special
emphasis on the role of zooplankton for water clarity.
Hydrobiologia 408/409: 217–231.
Khan FA and Ansari AA (2005) Eutrophication: an ecological
vision. Botanical Review 71: 449–482.
Lippert BE and Jameson DL (1964) Plant succession in temporary
ponds of the Willamette Valley, Oregon. American Midland
Naturalist 71: 181–197.
Lucassen ECHET, Smolders AJP, Lamers LPM and Roelofs
JGM (2005) Water table fluctuations and groundwater supply
are important in preventing phosphate-eutrophication in
sulphate-rich fens: consequences for wetland restoration. Plant
and Soil 269: 109–115.
Maberly SC and Madsen TV (1998) Affinity for CO2 in relation to
the ability of freshwater macrophytes to use HCO2 3 . Functional
Ecology 12: 99–106.
Middelboe AL and Markager S (1997) Depth limits and minimum
light requirements of freshwater macrophytes. Freshwater
Biology 37: 553–568.
Noon KF (1996) A model of created wetland primary succession.
Landscape and Urban Planning 34: 97–123.
Pezeshki SR (2001) Wetland plant responses to soil flood-
ing. Environmental and Experimental Botany 46: 299–
312.
Rascio N (2002) The underwater life of secondarily aquatic plants:
some problems and solutions. Critical Reviews in Plant Sciences
21: 401–427.
Ray AM, Rebertus AJ and Ray HL (2001) Macrophyte
succession in Minnesota beaver ponds. Canadian Journal of
Botany 79: 487–499.
Redding TE and Devito KJ (2006) Particle densities of wetland
soils in northern Alberta, Canada. Canadian Journal of Soil
Science 86: 57–60.
Sand-Jensen K and Jacobsen D (2002) Herbivory and growth in
terrestrial and aquatic populations of amphibious stream
plants. Freshwater Biology 47: 1475–1487.
Schaminee JHJ, van Kley JE and Ozinga WA (2002) The analysis
of long-term changes in plant communities: case studies from
the Netherlands. Phytocoenologia 32: 317–335.
Scheffer M, Hosper SH, Meijer ML, Moss B and Jeppesen E
(1993) Alternative equilibria in shallow lakes. Trends in Ecology
& Evolution 8: 175–279.
Schneider S (2007) Macrophyte trophic indicator values from a
European perspective. Limnologica 37: 281–289.
Schutten J, Dainty J and Davy AJ (2005) Root anchorage and its
significance for submerged plants in shallow lakes. Journal of
Ecology 93: 556–571.
Sculthorpe CD (1967) The Biology of Aquatic Vascular Plants.
London: Edward Arnold.
Smolders AJP, Lamers LPM, Lucassen ECHET, van der Velde G
and Roelofs JGM (2006) Internal eutrophication: how it works
and what to do about it – a review. Chemistry and Ecology 22:
93–111.

Smolders AJP, Vergeer LHT, van der Velde G and Roelofs JGM
(2003) Phenolic contents of submerged, emergent and floating
leaves of aquatic and semi-aquatic macrophyte species: why do
they differ? OIKOS 91: 307–310.
Sraj-Krzic N, Pongrac P, Klemenc M et al. (2006) Mycorrhizal
colonisation in plants from intermittent aquatic habitats.
Aquatic Botany 85: 333–338.
Tanaka N, Uehara K and Murata A (2004) Correlation between
pollen morphology and pollination mechanisms in the Hydro-
charitaceae. Journal of Plant Research 117: 265–276.
Tilman D (1982) Resource Competition and Community Structure.
Princeton: Princeton University Press.
Titus JE and Sullivan PG (2001) Heterophylly in the yellow
waterlily, Nuphar variegata (Nymphaeaceae): effects of [CO2],
natural sediment type, and water depth. American Journal of
Botany 88: 1469–1478.
Van der Valk AG (2005) Water level fluctuations in North
American prairie wetlands. Hydrobiologia 539: 171–188.
Vestergaard O and Sand-Jensen K (2000) Alkalinity and trophic
state regulate aquatic plant distribution in Danish lakes.
Aquatic Botany 67: 85–107.
Visser EJW, Colmer TD, Blom CWPM and Voesenek LACJ
(2000) Changes in growth, porosity, and radial oxygen loss from
adventitious roots of selected mono- and dycotyledonous
wetland species with contrasting types of aerenchyma. Plant,
Cell and Environment 23: 1237–1245.
Willby NJ, Abernethy VJ and Demars BOL (2000) Attribute-
based classification of European hydrophytes and its relation-
ship to habitat utilization. Freshwater Biology 43: 43–74.
Further Reading
Barendregt A and Bio AMF (2003) Relevant variables to predict
macrophyte communities in running waters. Ecological Model-
ling 160: 205–217.
Barrett SCH, Eckert CG and Husband BC (1993) Evolutionary
processes in aquatic plant populations. Aquatic Botany 44: 105–
145.
ENCYCLOPEDIA OF LIFE SCIENCES & 2009, John Wiley & Sons, Ltd. www.els.net
Macrophytes: Ecology of Aquatic Plants
Caraco N, Cole J, Findlay S and Wigand C (2006) Vascular plants
as engineers of oxygen in aquatic systems. BioScience 56: 219–
225.
Cook CDK (1990) Aquatic Plant Book. The Hague, The Nether-
lands: SPB Academic publishing.
Engelhardt KAM and Ritchie ME (2001) Effects of macrophyte
species richness on wetland ecosystem functioning and services.
Nature 441: 687–689.
Keeley JE (1998) CAM photosynthesis in submerged aquatic
plants. Botanical Review 64: 121–175.
Lacoul P and Freedman B (2006) Environmental influences on
aquatic plants in freshwater ecosystems. Environmental Review
14: 89–136.
Minorsky PV (2003) Heterophylly in aquatic plants. Plant
Physiology 133: 1671–1672.
Oertli B, Auderset Joye D, Castella E et al. (2005) PLOCH: a
standardized method for sampling and assessing the biodiver-
sity in ponds. Aquatic Conservation: Marine and Freshwater
Ecosystems 15: 665–679.
Pagano AM and Titus JE (2007) Submersed macrophyte growth
at low pH: carbon source influences response to dissolved
inorganic carbon enrichment. Freshwater Biology 52: 2412–
2420.
Puijalon S, Léna JP, Rivière N et al. (2008) Phenotypic plasticity in
response to mechanical stress: hydrodynamic performance and
fitness of 4 aquatic plant species. New Phytologist 177: 907–917.
Santamaria L (2002) Why are most aquatic plants widely
distributed? Dispersal, clonal growth and small-scale hetero-
geneity in a stressful environment. Acta Oecologica 23: 137–
154.
Sawada M, Viau AE and Gajewski K (2003) The biogeography of
aquatic macrophytes in North America since the Last Glacial
Maximum. Journal of Biogeography 30: 999–1017.
Wells CL and Pigliucci M (2000) Adaptive phenotypic plasticity:
the case of heterophylly in aquatic plants. Perspectives in Plant
Ecology, Evolution and Systematics 3: 1–18.
Wisheu IC and Keddy PA (1992) Competition and centrifugal
organization of plant communities: theory and tests. Journal of
Vegetation Science 3: 147–156.
9