Parasitic plant

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Cuscuta, a stem holoparasite, on an Acacia tree in Pakistan Cuscuta parasite plant.JPG
Cuscuta , a stem holoparasite, on an Acacia tree in Pakistan

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 in the air or soil given off by host shoots or roots, respectively. About 4,500 species of parasitic plants in approximately 20 families of flowering plants are known. [5] [6]

Contents

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 families have only a few representatives. One example is the 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]

  1     a    Obligate An obligate parasite cannot complete its life cycle without a host.
b  Facultative  A facultative parasite can complete its life cycle independent of a host.
2aStemA stem parasite attaches to the host stem.
bRootA root parasite attaches to the host root.
3aHemi-A hemiparasitic plant lives as a parasite under natural conditions, but remains photosynthetic 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.
bHolo-A holoparasitic plant derives all of its fixed carbon from the host plant. Commonly lacking chlorophyll, holoparasites are often colors that are not green.
Mistletoe, an obligate stem hemiparasite Mistletoe, coming soon to a market near you - geograph.org.uk - 1585249.jpg
Mistletoe, an obligate stem hemiparasite

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

Holoparasites are always obligate so only two terms are needed, e.g.

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

Striga witchweeds (white, center, attached to roots of the host) are economically important pests of the crop plants that they parasitize. Striga root connections.jpg
Striga witchweeds (white, center, attached to roots of the host) are economically important pests of the crop plants that they parasitize.

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 whether or not the parasites are root parasites or stem parasites. Most parasitic plants need to germinate near their host plants because their seeds are limited in the number of resources necessary to survive without 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 6 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]

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]

Newly emergent snow plant (Sarcodes sanguinea), a flowering plant parasitic on mycorrhizal fungi Sarcodes sanguinea 08080.jpg
Newly emergent snow plant ( Sarcodes sanguinea ), a flowering plant parasitic on mycorrhizal fungi

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 gives these plants the ability to become associated with many of the other plants around them. They are termed myco-heterotrophs. 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]

Related Research Articles

<i>Cuscuta</i> Genus of parasitic plants

Cuscuta, commonly known as dodder or amarbel, is a genus of over 201 species of yellow, orange, or red parasitic plants. Formerly treated as the only genus in the family Cuscutaceae, it now is accepted as belonging in the morning glory family, Convolvulaceae, on the basis of the work of the Angiosperm Phylogeny Group. The genus is found throughout the temperate and tropical regions of the world, with the greatest species diversity in subtropical and tropical regions; the genus becomes rare in cool temperate climates, with only four species native to northern Europe.

<span class="mw-page-title-main">Mistletoe</span> Common name for various parasitic plants that grow on trees and shrubs

Mistletoe is the common name for obligate hemiparasitic plants in the order Santalales. They are attached to their host tree or shrub by a structure called the haustorium, through which they extract water and nutrients from the host plant.

<span class="mw-page-title-main">Loranthaceae</span> Family of mistletoes

Loranthaceae, commonly known as the showy mistletoes, is a family of flowering plants. It consists of about 75 genera and 1,000 species of woody plants, many of them hemiparasites. The three terrestrial species are Nuytsia floribunda, Atkinsonia ligustrina, and Gaiadendron punctatum Loranthaceae are primarily xylem parasites, but their haustoria may sometimes tap the phloem, while Tristerix aphyllus is almost holoparasitic. For a more complete description of the Australian Loranthaceae, see Flora of Australia online., for the Malesian Loranthaceae see Flora of Malesia.

<span class="mw-page-title-main">Orobanchaceae</span> Family of flowering plants known as broomrapes

Orobanchaceae, the broomrapes, is a family of mostly parasitic plants of the order Lamiales, with about 90 genera and more than 2000 species. Many of these genera were formerly included in the family Scrophulariaceae sensu lato. With its new circumscription, Orobanchaceae forms a distinct, monophyletic family. From a phylogenetic perspective, it is defined as the largest crown clade containing Orobanche major and relatives, but neither Paulownia tomentosa nor Phryma leptostachya nor Mazus japonicus.

<i>Striga</i> Genus of flowering plants belonging to the broomrape family

Striga, commonly known as witchweed, is a genus of parasitic plants that occur naturally in parts of Africa, Asia, and Australia. It is currently classified in the family Orobanchaceae, although older classifications place it in the Scrophulariaceae. Some species are serious pathogens of cereal crops, with the greatest effects being in savanna agriculture in Africa. It also causes considerable crop losses in other regions, including other tropical and subtropical crops in its native range and in the Americas. The generic name derives from Latin strī̆ga, "witch".

<i>Cassytha</i> Genus of flowering plants

Cassytha is a genus of some two dozen species of obligately parasitic vines in the family Lauraceae. Superficially, and in some aspects of their ecology, they closely resemble plants in the unrelated genus Cuscuta, the dodders. When fruit and flowers are absent in the field, the physical resemblance is so close that few people without technical training can discern the difference. In this respect and in their ecology the two genera present a spectacular example of convergent evolution. Nonetheless, Nickrent comments that "Cassytha is uneqivocally assigned to Lauraceae based on (both) morphological and molecular data." In its divergence from habits typical of the Lauraceae, Cassytha also presents examples of mosaic evolution

<i>Phoradendron</i> Genus of mistletoes

Phoradendron is a genus of mistletoe, native to warm temperate and tropical regions of the Americas. The center of diversity is the Amazon rainforest. Phoradendron is the largest genus of mistletoe in the Americas, and possibly the largest genus of mistletoes in the world. Traditionally, the genus has been placed in the family Viscaceae, but recent genetic research acknowledged by the Angiosperm Phylogeny Group shows this family to be correctly placed within a larger circumscription of the sandalwood family, Santalaceae.

<i>Orobanche aegyptiaca</i> Species of flowering plant

Orobanche aegyptiaca, the Egyptian broomrape, is a plant which is an obligate holoparasite from the family Orobanchaceae with a complex lifecycle. This parasite is most common in the Middle East and has a wide host range including many economically important crops.

<i>Triphysaria</i> Genus of flowering plants belonging to the broomrape family

Triphysaria is a genus of five plants in the family Orobanchaceae which are known generally as owl's-clovers. This genus is closely related to the genera Castilleja and Orthocarpus. Triphysaria are native to western North America, including a species endemic to California.

<i>Cuscuta campestris</i> Species of flowering plant

Cuscuta campestris, with the common names field dodder, golden dodder, large-seeded alfalfa dodder, yellow dodder and prairie dodder, is a parasitic plant which belongs to the family Convolvulaceae. It was formerly classified in the family Cuscutaceae.

<i>Orobanche minor</i> Species of flowering plant

Orobanche minor, the hellroot, common broomrape, lesser broomrape, small broomrape or clover broomrape, is a holoparasitic flowering plant belonging to the family Orobanchaceae. It is one of about 150 non-photosynthetic plants in the genus Orobanche that parasitize autotrophic plants.

<i>Striga hermonthica</i> Species of flowering plant

Striga hermonthica, commonly known as purple witchweed or giant witchweed, is a hemiparasitic plant that belongs to the family Orobanchaceae. It is devastating to major crops such as sorghum and rice. In sub-Saharan Africa, apart from sorghum and rice, it also infests maize, pearl millet, and sugar cane.

<i>Cassytha pubescens</i> Species of plant

Cassytha pubescens is a native Australian hemiparasitic vine species, in the Laurel family. Common names for the species include devils twine, dodder-laurel, spilled devil's twine, snotty gobble or downy dodder-laurel. It is a widespread and common species in south eastern Australia. The species was first formally described in 1810 by the Scottish botanist Robert Brown in Prodromus Flora Novae Hollandiae et Insulae Van Diemen. Leaves are reduced to scales and photosynthesis is achieved through chlorophyll contained in the plants stems. Stems are between 0.5mm and 1.5mm in diameter and the haustoria are between 2 and 3 mm long.

<i>Phoradendron tomentosum</i> Species of mistletoe

Phoradendron tomentosum, the leafy mistletoe, hairy mistletoe or Christmas mistletoe, is a plant parasite. It is characterized by its larger leaves and smaller berries than dwarf mistletoe. Leafy mistletoe seldom kill but they do rob their hosts of moisture and some minerals, causing stress during drought and reducing crop productions on fruit and nut trees. Leafy mistletoe has the ability to photosynthesize on its own but it relies on other plants in order to obtain its nutrients. It attaches itself to a tree and then grows haustoria, in order to get the food and water it needs.

<i>Arceuthobium pusillum</i> Species of dwarf mistletoe

Arceuthobium pusillum is a perennial, obligate parasitic plant in the sandalwood family. Its common names include Dwarf mistletoe or Eastern dwarf mistletoe. It is one of the most widespread dwarf mistletoes within its range which covers the eastern United States and Canada, from Saskatchewan to Nova Scotia and New Jersey. The species name "pusillum" derives from Latin "pusillus", meaning very small.

<i>Orobanche hederae</i> Species of flowering plant

Orobanche hederae, the ivy broomrape, is, like other members of the genus Orobanche, a parasitic plant without chlorophyll, and thus totally dependent on its host, which is ivy. It grows to 60 cm (2 ft), with stems in shades of brown and purple, sometimes yellow. The flowers are 10–22 mm (0.4–0.9 in) long, cream in colour with reddish-purple veins.

<i>Cuscuta compacta</i> Species of flowering plant

Cuscuta compacta, the compact dodder, is a parasitic plant that specializes on woody plants. This species is distributed across the Eastern and Midwestern USA, Eastern Canada, and Mexico.

<span class="mw-page-title-main">Strigolactone</span> Group of chemical compounds

Strigolactones are a group of chemical compounds produced by roots of plants. Due to their mechanism of action, these molecules have been classified as plant hormones or phytohormones. So far, strigolactones have been identified to be responsible for three different physiological processes: First, they promote the germination of parasitic organisms that grow in the host plant's roots, such as Strigalutea and other plants of the genus Striga. Second, strigolactones are fundamental for the recognition of the plant by symbiotic fungi, especially arbuscular mycorrhizal fungi, because they establish a mutualistic association with these plants, and provide phosphate and other soil nutrients. Third, strigolactones have been identified as branching inhibition hormones in plants; when present, these compounds prevent excess bud growing in stem terminals, stopping the branching mechanism in plants.

<i>Cuscuta gronovii</i> Species of flowering plant

Cuscuta gronovii is a yellow vine that grows as a parasite off other plants. It is a dicot.

Steven Runo is a Kenyan researcher who is a professor at Kenyatta University. Runo has extensively studied the pathogens of African cereal crops, including Striga. He was awarded the 2020 Royal Society Africa Prize.

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  29. ↑ Těšitel, Jakub; Li, Ai-Rong; Knotková, Kateřina; McLellan, Richard; Bandaranayake, Pradeepa C G; Watson, David M (April 2021). "The bright side of parasitic plants: what are they good for?". Plant Physiology. 185 (4): 1309–1324. doi:10.1093/plphys/kiaa069. PMC   8133642 . PMID   33793868 – via Oxford Academic.
  30. ↑ O'Neill, Alexander; Rana, Santosh (2017-07-16). "An ethnobotanical analysis of parasitic plants (Parijibi) in the Nepal Himalaya". Journal of Ethnobiology and Ethnomedicine. 12 (14): 14. doi: 10.1186/s13002-016-0086-y . PMC   4765049 . PMID   26912113.
  31. ↑ Judd, Walter S., Christopher Campbell, and Elizabeth A. Kellogg. Plant Systematics: A Phylogenetic Approach. Sunderland, MA: Sinauer Associates, 2008. Print.
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