Parasitic plant

A parasitic plant is a plant that derives some or all of

its nutritional requirements from another living plant.
They make up about 1% of angiosperms and are found
in almost every biome. All parasitic plants develop a
specialized organ called the haustorium, which
penetrates the host plant, connecting them to the host
vasculature – either the xylem, phloem, or both.[1] For
example, plants like Striga or Rhinanthus connect only
to the xylem, via xylem bridges (xylem-feeding).
Alternately, plants like Cuscuta and some members of
Orobanche connect to both the xylem and phloem of
the host.[1][2][3] This provides them with the ability to
extract resources from the host. These resources can
include water, nitrogen, carbon and/or sugars.[4]
Parasitic plants are classified depending on the location
where the parasitic plant latches onto the host (root or
stem), the amount of nutrients it requires, and their
photosynthetic capability.[5] Some parasitic plants can
locate their host plants by detecting volatile chemicals
Cuscuta, a stem holoparasite, on an Acacia tree
in the air or soil given off by host shoots or roots,
in Pakistan
respectively. About 4,500 species of parasitic plants in
approximately 20 families of flowering plants are
known.[5][6]

There is a wide range of effects that may occur to a host plant due
to the presence of a parasitic plant. Often there is a pattern of
stunted growth in hosts especially in hemi-parasitic cases, but may
also result in higher mortality rates in host plant species following
introduction of larger parasitic plant populations.[7]

Classification
Parasitic plants occur in multiple plant families, indicating that the
evolution is polyphyletic. Some families consist mostly of
parasitic representatives such as Balanophoraceae, while other Rafflesia arnoldii, another type of
families have only a few representatives. One example is the parasitic plant
North American Monotropa uniflora (Indian pipe or corpse plant)
which is a member of the heath family, Ericaceae, better known
for its member blueberries, cranberries, and rhododendrons.

Parasitic plants are characterized as follows:[5]

 a Obligate An obligate parasite cannot complete its life cycle without a host.
1
b Facultative A facultative parasite can complete its life cycle independent of a host.

a Stem A stem parasite attaches to the host stem.

2
b Root A root parasite attaches to the host root.

A hemiparasitic plant lives as a parasite under natural conditions, but remains photosynthetic
a Hemi- to at least some degree. Hemiparasites may obtain only water and mineral nutrients from the
host plant, or many also obtain a part of their organic nutrients from the host.
3
A holoparasitic plant derives all of its fixed carbon from the host plant. Commonly lacking
b Holo-
chlorophyll, holoparasites are often colors that are not green.

For hemiparasites, one from each of the three sets of terms can be
applied to the same species, e.g.

Nuytsia floribunda (Western Australian Christmas tree) is

an obligate root hemiparasite.
Rhinanthus (e.g. Yellow rattle) is a facultative root
hemiparasite.
Mistletoe is an obligate stem hemiparasite.
Holoparasites are always obligate so only two terms are needed,
Mistletoe, an obligate stem
e.g.
hemiparasite

Dodder is a stem holoparasite.

Hydnora spp. are root holoparasites.
Plants usually considered holoparasites include broomrape, dodder, Rafflesia, and the Hydnoraceae.
Plants usually considered hemiparasites include Castilleja, mistletoe, Western Australian Christmas tree,
and yellow rattle.

Evolution of parasitism
Parasitic behavior evolved in angiosperms roughly 12-13 times independently, a classic example of
convergent evolution. Roughly 1% of all angiosperm species are parasitic, with a large degree of host
dependence. The taxonomic family Orobanchaceae (encompassing the genera Triphysaria, Striga, and
Orobanche) is the only family that contains both holoparasitic and hemiparasitic species, making it a
model group for studying the evolutionary rise of parasitism. The remaining groups contain only
hemiparasites or holoparasites.[8]

The evolutionary event which gave rise to parasitism in plants was the development of haustoria. The
first, most ancestral, haustoria are thought to be similar to that of the facultative hemiparasites within
Triphysaria, lateral haustoria develop along the surface of the roots in these species. Later evolution led
to the development of terminal or primary haustoria at the tip of the juvenile radicle, seen in obligate
hemiparasitic species within Striga. Lastly, holoparasitic plants, always forms of obligate parasites,
evolved over the loss of photosynthesis, seen in the genus Orobanche.[8] The most specialized forms of
holoparasitic plants are the four families Rafflesiaceae, Cytinaceae, Mitrastemonaceae and
Apodanthaceae, lineages which independently has evolved further into endoparasites that, except for the
flowers, spend their entire life cycle within the tissue of their host.[9]
To maximize resources, many parasitic plants have evolved 'self-
incompatibility', to avoid parasitizing themselves. Others such as
Triphysaria usually avoid parasitizing other members of their species, but
some parasitic plants have no such limits.[8] The albino redwood is a
mutant Sequoia sempervirens that produces no chlorophyll; they live on
sugars from neighbouring trees, usually the parent tree from which they
have grown (via a somatic mutation).[10][11][12]

Seed germination
Parasitic plants germinate in several methods. These can either be
chemical or mechanical and the means used by seeds often depends on Striga witchweeds (white,
whether or not the parasites are root parasites or stem parasites. Most center, attached to roots of
parasitic plants need to germinate near their host plants because their the host) are economically
seeds are limited in the number of resources necessary to survive without important pests of the crop
plants that they parasitize.
nutrients from their host plants. Resources are limited due in part to the
fact that most parasitic plants are not able to use autotrophic nutrition to
establish the early stages of seeding.[13][14]

Root parasitic plant seeds tend to use chemical cues for germination. For germination to occur, seeds
need to be quite close to the host plant.[13][14] For example, the seeds of witchweed (Striga asiatica) need
to be within 3 to 4 millimeters (mm) of its host to receive chemical signals in the soil to trigger
germination. This range is important because Striga asiatica will only grow about 4 mm after
germination.[13] Chemical compound cues sensed by parasitic plant seeds are from host plant root
exudates that are leached nearby from the host's root system into the surrounding soil. These chemical
cues are a variety of compounds that are unstable and rapidly degraded in soil and are present within a
radius of a few meters of the plant exuding them. Parasitic plants germinate and follow a concentration
gradient of these compounds in the soil toward the host plants if close enough. These compounds are
called strigolactones. Strigolactone stimulates ethylene biosynthesis in seeds causing them to
germinate.[13][14]

There are a variety of chemical germination stimulants. Strigol was the first of the germination stimulants
to be isolated. It was isolated from a non-host cotton plant and has been found in true host plants such as
corn and millets. The stimulants are usually plant-specific, examples of other germination stimulants
include sorgolactone from sorghum, Orobanche and electoral from red clover, and 5-deoxystrigol from
Lotus japonicus. Strigolactones are apocarotenoids that are produced via the carotenoid pathway of
plants. Strigolactones and mycorrhizal fungi have a relationship in which Strigolactone also cues the
growth of mycorrhizal fungus.[14][15]

Stem parasitic plants, unlike most root parasites, germinate using the resources inside their endosperms
and can survive for some time. For example, the dodders (Cuscuta spp.) drop their seeds to the ground.
These may remain dormant for up to five years before they find a host plant. Using the resources in the
seed endosperm, the dodder can germinate. Once germinated, the plant has six days to find and establish
a connection with its host plant before its resources are exhausted.[13] Dodder seeds germinate above
ground, then the plant sends out stems in search of its host plant reaching up to 6 cm before it dies. It is
believed that the plant uses two methods of finding a host. The stem detects its host plant's scent and
orients itself in that direction. Scientists used volatiles from tomato plants (α-pinene, β-myrcene, and β-
phellandrene) to test the reaction of C. pentagona and found that the stem orients itself in the direction of
the odor.[14] Some studies suggest that by using light reflecting from nearby plants dodders can select
hosts with higher sugar because of the levels of chlorophyll in the leaves.[16] Once the dodder finds its
host, it wraps itself around the host plant's stem. Using adventitious roots, the dodder taps into the host
plant's stem with a haustorium, an absorptive organ within the host plant vascular tissue. Dodder makes
several of these connections with the host as it moves up the plant.[13][14][16]

Seed dispersal
There are several methods of seed dispersal, but all the strategies aim to put the seed in direct contact
with, or within a critical distance of, the host.

1. The Cuscuta seedling can live for 3–7 days and extend out 35 cm in search of the host
before it dies. This is because the Cuscuta seed is large and has stored nutrients to sustain
its life. This is also useful for seeds that get digested by animals and are excreted.[5]
2. Mistletoe use a sticky seed for dispersal. The seed sticks to nearby animals and birds and
then comes into direct contact with the host.[5]
3. Arceuthobium seeds have a similarly sticky seed as the mistletoe but they do not rely on
animals and birds, they mainly disperse by fruit explosiveness. Once the seed makes
contact with the host, rainwater can help position the seed in a suitable position.[5]
4. Some seeds detect and respond to chemical stimulations produced in the host's roots and
start to grow towards the host.[5]

Obstacles to host attachment

A parasitic plant has many obstacles to overcome to attach to a host. Distance from the host and stored
nutrients are some of the problems, and the host's defenses are an obstacle to overcome. The first hurdle
is penetrating the host since the host has systems to reinforce the cell wall by protein cross-linking so that
it stops the parasitic progress at the cortex of the host's roots. The second hurdle is the host's ability to
secrete germination inhibitors. This prevents germination of the parasitic seed. The third hurdle is the
host's ability to create a toxic environment at the location where the parasitic plant attaches. The host
secretes phenolic compounds into the apoplast. This creates a toxic environment for the parasitic plant,
eventually killing it. The fourth hurdle is the host's ability to ruin the tubercle using gums and gels or
injecting toxins into the tubercle.[17]

Host range
Some parasitic plants are generalists and parasitize many different species, even several different species
at once.[18] Dodder (Cuscuta spp.) and red rattle (Odontites vernus) are generalist parasites. Other
parasitic plants are specialists that parasitize a few or just one species. Beech drops (Epifagus virginiana)
is a root holoparasite only on American beech (Fagus grandifolia). Rafflesia is a holoparasite on the vine
Tetrastigma. Plants such as Pterospora become parasites of mycorrhizal fungi. There is evidence that
parasites also practice self-discrimination, species of Triphysaria experience reduced haustorium
development in the presence of other Triphysaria. The mechanism for self-discrimination in parasites is
not yet known.[8]

Aquatic parasitic plants

Parasitism also evolved within aquatic species of plants and algae. Parasitic marine plants are described
as benthic, meaning that they are sedentary or attached to another structure. Plants and algae that grow on
the host plant, using it as an attachment point are given the designation epiphytic (epilithic is the name
given to plants/algae that use rocks or boulders for attachment), while not necessarily parasitic, some
species occur in high correlation with a certain host species, suggesting that they rely on the host plant in
some way or another. In contrast, endophytic plants and algae grow inside their host plant, these have a
wide range of host dependence from obligate holoparasites to facultative hemiparasites.[19]

Marine parasites occur as a higher proportion of marine flora in temperate rather than tropical waters.
While no full explanation for this is available, many of the potential host plants such as kelp and other
macroscopic brown algae are generally restricted to temperate areas. Roughly 75% of parasitic red algae
infect hosts in the same taxonomic family as themselves, these are given the designation
adelphoparasites. Other marine parasites, deemed endozoic, are parasites of marine invertebrates
(mollusks, flatworms, sponges) and can be either holoparasitic or hemiparasitic, some retaining the ability
to photosynthesize after infection. These are the only parasitic plants that parasitize animal hosts.[19]

Importance
Species within Orobanchaceae are some of the most economically destructive species on Earth. Species
of Striga alone are estimated to cost billions of dollars a year in crop yield loss annually, infesting over 50
million hectares of cultivated land within sub-Saharan Africa alone. Striga can infest both grasses and
grains, including corn, rice and sorghum, some of the most important food crops. Orobanche also
threatens a wide range of important crops, including peas, chickpeas, tomatoes, carrots, lettuce,[20] and
varieties of the genus Brassica (e.g. cabbage and broccoli). Yield loss from Orobanche can reach 100%
and has caused farmers in some regions of the world to abandon certain staple crops and begin importing
others as an alternative. Much research has been devoted to the control of Orobanche and Striga species,
which are even more devastating in developing areas of the world, though no method has been found to
be entirely successful.[8]

Mistletoes cause economic damage to forests and ornamental trees.

Rafflesia arnoldii produces the world's largest flowers at about one meter in diameter. It is a
tourist attraction in its native habitat.
Sandalwood trees (Santalum species) have many important cultural uses and their fragrant
oils have high commercial value.
Indian paintbrush (Castilleja linariaefolia) is the state flower of Wyoming.
The oak mistletoe (Phoradendron serotinum) is the floral emblem of Oklahoma.
A few other parasitic plants are occasionally cultivated for their attractive flowers, such as
Nuytsia and broomrape.
Parasitic plants are important in research, especially on the loss of photosynthesis and the
 co-dependency of functional, genetic and lifestyle changes.[21][22][23][24]
A few dozen parasitic plants have occasionally been used as food by people.[25]
Western Australian Christmas tree (Nuytsia floribunda) sometimes damages underground
cables. It mistakes the cables for host roots and tries to parasitize them using its
sclerenchymatic guillotine.[26]
Some parasitic plants are destructive while some have positive influences in their communities. Some
parasitic plants damage invasive species more than native species. This results in the reduced damage of
invasive species in the community.[27] Parasitic plants are major shapers of their community, affecting not
just the host species but indirectly affecting others. Competition amongst host species will change due to
the parasitic plant.[28] Plant parasitism have been shown to keep invasive species under control and
become keystone species in an ecosystem.[29]

In many regions, including the Nepal Eastern Himalayas, parasitic

plants are used for medicinal and ritual purposes.[30]

Plants parasitic on fungi

About 400 species of flowering plants, plus one gymnosperm
(Parasitaxus usta) and one bryophyte (the liverwort Aneura
mirabilis), are parasitic on mycorrhizal fungi. This effectively
Newly emergent snow plant
gives these plants the ability to become associated with many of (Sarcodes sanguinea), a flowering
the other plants around them. They are termed myco-heterotrophs. plant parasitic on mycorrhizal fungi
Some myco-heterotrophs are Indian pipe (Monotropa uniflora),
snow plant (Sarcodes sanguinea), underground orchid
(Rhizanthella gardneri), bird's nest orchid (Neottia nidus-avis), and sugarstick (Allotropa virgata). Within
the taxonomic family Ericaceae, known for extensive mycorrhizal relationships, there are the
Monotropoids. The Monotropoids include the genera Monotropa, Monotropsis, and Pterospora among
others. Myco-heterotrophic behavior is commonly accompanied by the loss of chlorophyll.[31]

See also
Living stump

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External links
Media related to Parasitic plants at Wikimedia Commons
The International Parasitic Plant Society (https://www.parasiticplants.org/)

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