Seagrass

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Zostera marina is the most abundant seagrass species in the Northern Hemisphere

Seagrasses are the only flowering plants which grow in marine environments. There are about 60 species of fully marine seagrasses which belong to four families (Posidoniaceae, Zosteraceae, Hydrocharitaceae and Cymodoceaceae), all in the order Alismatales (in the class of monocotyledons).[1] Seagrasses evolved from terrestrial plants which recolonised the ocean 70 to 100 million years ago.

The name seagrass stems from the many species with long and narrow leaves, which grow by rhizome extension and often spread across large "meadows" resembling grassland; many species superficially resemble terrestrial grasses of the family Poaceae.

Like all autotrophic plants, seagrasses photosynthesize, in the submerged photic zone, and most occur in shallow and sheltered coastal waters anchored in sand or mud bottoms. Most species undergo submarine pollination and complete their life cycle underwater.

Seagrasses form dense underwater seagrass meadows which are among the most productive ecosystems in the world. They function as important carbon sinks and provide habitats and food for a diversity of marine life comparable to that of coral reefs.

Overview[edit]

Seagrasses are a paraphyletic group of marine angiosperms which evolved three to four times from land plants back to the sea. The following characteristics can be used to define a seagrass species. It lives in an estuarine or in the marine environment, and nowhere else. The pollination takes place underwater with specialized pollen. The seeds which are dispersed by both biotic and abiotic agents are produced underwater.[2] The seagrass species have specialized leaves with a reduced cuticle, an epidermis which lacks stomata and is the main photosynthetic tissue. The rhizome or underground stem is important in anchoring. The roots can live in an anoxic environment and depend on oxygen transport from the leaves and rhizomes but are also important in the nutrient transfer processes.[3][2]

Seagrasses profoundly influence the physical, chemical, and biological environments of coastal waters.[2] Though seagrasses provide invaluable ecosystem services by acting as breeding and nursery ground for a variety of organisms and promote commercial fisheries, many aspects of their physiology are not well investigated. Several studies have indicated that seagrass habitat is declining worldwide.[4][5] Ten seagrass species are at elevated risk of extinction (14% of all seagrass species) with three species qualifying as endangered. Seagrass loss and degradation of seagrass biodiversity will have serious repercussions for marine biodiversity and the human population that depends upon the resources and ecosystem services that seagrasses provide.[6][2]

Evolution[edit]

Evolution of seagrass
Showing the progression onto land from marine origins, the diversification of land plants and the subsequent return to the sea by the seagrasses.

Terrestrial plants evolved perhaps as early as 450 million years ago from a group of green algae.[7] Seagrasses then evolved from terrestrial plants which migrated back into the ocean.[8][9] Between about 70 million and 100 million years ago, the three independent seagrass lineages (Hydrocharitaceae, Cymodoceaceae complex, and Zosteraceae) evolved from a single lineage of monocotyledonous flowering plants.[10]

Other plants that colonised the sea, such as salt marsh plants, mangroves, and marine algae, have more diverse evolutionary lineages. In spite of their low species diversity, seagrasses have succeeded in colonising the continental shelves of all continents except Antarctica.[11]

Around 140 million years ago, seagrasses evolved from early monocotyledonous land plants, which succeeded in conquering the marine environment. Today, they are a polyphyletic group of marine angiosperms with around 60 species in five families (Zosteraceae, Hydrocharitaceae, Posidoniaceae, Cymodoceaceae, and Ruppiaceae), which belong to the order Alismatales according to the Angiosperm Phylogeny Group IV System.[12] The genus Ruppia, which occurs in brackish water, is not regarded as a "real" seagrass by all authors and has been shifted to the Cymodoceaceae by some authors.[13] The APG IV system and The Plant List Webpage[14] do not share this family assignment.[15]

Seagrasses form important coastal ecosystems.[16] The worldwide endangering of these sea meadows, which provide food and habitat for many marine species, prompts the need for protection and understanding of these valuable resources. Recent sequencing of the genomes of Zostera marina and Zostera muelleri has given better understanding angiosperm adaption to the sea.[17][18] During the evolutionary step back to the ocean, different genes have been lost (e.g., stomatal genes) or have been reduced (e.g., genes involved in the synthesis of terpenoids) and others have been regained, such as in genes involved in sulfation.[18][15]

Genome information has shown further that adaption to the marine habitat was accomplished by radical changes in cell wall composition.[17][18] However the cell walls of seagrasses are not well understood. In addition to the ancestral traits of land plants one would expect habitat-driven adaption processs to the new environment characterized by multiple abiotic (high amounts of salt) and biotic (different seagrass grazers and bacterial colonization) stressors.[15] The cell walls of seagrasses seem intricate combinations of features known from both angiosperm land plants and marine macroalgae with new structural elements.[15]

Taxonomy[edit]

Family Image Genera Description
Zosteraceae The family Zosteraceae, also known as the seagrass family, includes two genera containing 14 marine species. It is found in temperate and subtropical coastal waters, with the highest diversity located around Korea and Japan.

Species subtotal:  

Tectura palacea 3.jpg
Phyllospadix 6 species  
Zostera.jpg
Zostera 16 species  
Hydrocharitaceae The family Hydrocharitaceae, also known as tape-grasses, include Canadian waterweed and frogbit. The family includes both fresh and marine aquatics, although of the sixteen genera currently recognised, only three are marine.[19] They are found throughout the world in a wide variety of habitats, but are primarily tropical.

Species subtotal:  

Enhalus acoroides01.jpg
Enhalus 1 species  
Johnsons seagrass bed.jpg
Halophila 19 species  
Thalassia hemprichii.jpg
Thalassia 2 species  
Posidoniaceae The family Posidoniaceae contains a single genus with two to nine marine species found in the seas of the Mediterranean and around the south coast of Australia.

Species subtotal: 2 to 9  

Posidonia 2 Alberto Romeo.jpg
Posidonia 2 to 9 species  
Cymodoceaceae The family Cymodoceaceae, also known as manatee-grass, includes only marine species.[20] Some taxonomists do not recognize this family.

Species subtotal:  

Amphibolis griffithii 34184967.jpg
Amphibolis 2 species  
Cymodocea.JPG
Cymodocea 4 species  
Halodule wrightii.jpg
Halodule 6 species  
Syringodium isoetifolium et Acropora sp..jpg
Syringodium 2 species  
Thalassodendron ciliatum.jpg
Thalassodendron 3 species  
Total species:   

Sexual recruitment[edit]

Seeds from Posidonia oceanica[21]
(A) Newly released seeds                 (B) One-week-old
           inside a fruit                                       seeds
FP: fruit pericarp; NRS: newly released seeds; WS: 1-week-old seeds; H: adhesive hairs; S: seed; R1: primary root; Rh: rhizome; L: leaves
The sexual recruitment stages of Posidonia oceanica[21]
Dispersion, adhesion and settlement

Seagrass populations are currently threatened by a variety of anthropogenic stressors.[22][5] The ability of seagrasses to cope with environmental perturbations depends, to some extent, on genetic variability, which is obtained through sexual recruitment.[23][24][25] By forming new individuals, seagrasses increase their genetic diversity and thus their ability to colonise new areas and to adapt to environmental changes.[26][27][28][29][30][21][excessive citations]

Seagrasses have contrasting colonisation strategies.[31] Some seagrasses form seed banks of small seeds with hard pericarps that can remain in the dormancy stage for several months. These seagrasses are generally short-lived and can recover quickly from disturbances by not germinating far away from parent meadows (e.g., Halophila sp., Halodule sp., Cymodocea sp., Zostera sp. and Heterozostera sp.[31][32] In contrast, other seagrasses form dispersal propagules. This strategy is typical of long-lived seagrasses that can form buoyant fruits with inner large non-dormant seeds, such as the genera Posidonia sp., Enhalus sp. and Thalassia sp.[31][33] Accordingly, the seeds of long-lived seagrasses have a large dispersal capacity compared to the seeds of the short-lived type,[34] which permits the evolution of species beyond unfavourable light conditions by the seedling development of parent meadows.[21]

The seagrass Posidonia oceanica (L.) Delile is one of the oldest and largest species on Earth. An individual can form meadows measuring nearly 15 km wide and can be as much as 100,000 years old.[35] P. oceanica meadows play important roles in the maintenance of the geomorphology of Mediterranean coasts, which, among others, makes this seagrass a priority habitat of conservation.[36] Currently, the flowering and recruitment of P. oceanica seems to be more frequent than that expected in the past.[37][38][39][40][41] Further, this seagrass has singular adaptations to increase its survival during recruitment. The large amounts of nutrient reserves contained in the seeds of this seagrass support shoot and root growth, even up to the first year of seedling development.[35] In the first months of germination, when leaf development is scarce, P. oceanica seeds perform photosynthetic activity, which increases their photosynthetic rates and thus maximises seedling establishment success.[42][43] Seedlings also show high morphology plasticity during their root system development [44][45] by forming adhesive root hairs to help anchor themselves to rocky sediments.[37][46][47] However, many factors about P. oceanica sexual recruitment remain unknown, such as when photosynthesis in seeds is active or how seeds can remain anchored to and persist on substrate until their root systems have completely developed.[21]

Intertidal and subtidal seagrasses[edit]

Morphological and photoacclimatory responses
of intertidal and subtidal Zostera marina eelgrass [48]

Seagrasses occurring in the intertidal and subtidal zones are exposed to highly variable environmental conditions due to tidal changes.[49][50] Subtidal seagrasses are more frequently exposed to lower light conditions, driven by plethora of natural and human-caused influences that reduce light penetration by increasing the density of suspended opaque materials. Subtidal light conditions can be estimated, with high accuracy, using artificial intelligence, enabling more rapid mitigation than was available using in situ techniques.[51] Seagrasses in the intertidal zone are regularly exposed to air and consequently experience extreme high and low temperatures, high photoinhibitory irradiance, and desiccation stress relative to subtidal seagrass.[50][52][53] Such extreme temperatures can lead to significant seagrass dieback when seagrasses are exposed to air during low tide.[54][55][56] Desiccation stress during low tide has been considered the primary factor limiting seagrass distribution at the upper intertidal zone.[57] Seagrasses residing the intertidal zone are usually smaller than those in the subtidal zone to minimize the effects of emergence stress.[58][55] Intertidal seagrasses also show light-dependent responses, such as decreased photosynthetic efficiency and increased photoprotection during periods of high irradiance and air exposure.[59][60]

Zostera marina seedling [61]

In contrast, seagrasses in the subtidal zone adapt to reduced light conditions caused by light attenuation and scattering due to the overlaying water column and suspended particles.[62][63] Seagrasses in the deep subtidal zone generally have longer leaves and wider leaf blades than those in the shallow subtidal or intertidal zone, which allows more photosynthesis, in turn resulting in greater growth.[53] Seagrasses also respond to reduced light conditions by increasing chlorophyll content and decreasing the chlorophyll a/b ratio to enhance light absorption efficiency by using the abundant wavelengths efficiently.[64][65][66] As seagrasses in the intertidal and subtidal zones are under highly different light conditions, they exhibit distinctly different photoacclimatory responses to maximize photosynthetic activity and photoprotection from excess irradiance.

Seagrasses assimilate large amounts of inorganic carbon to achieve high level production.[67][68] Marine macrophytes, including seagrass, use both CO2 and HCO
3
(bicarbonate) for photosynthetic carbon reduction.[69][70][71] Despite air exposure during low tide, seagrasses in the intertidal zone can continue to photosynthesize utilizing CO2 in the air.[72] Thus, the composition of inorganic carbon sources for seagrass photosynthesis probably varies between intertidal and subtidal plants. Because stable carbon isotope ratios of plant tissues change based on the inorganic carbon sources for photosynthesis,[73][74] seagrasses in the intertidal and subtidal zones may have different stable carbon isotope ratio ranges.

Seagrass microbiome[edit]

Processes within the seagrass holobiont
The most important interconnected processes within the seagrass holobiont are related to processes in the carbon-, nitrogen- and sulfur cycles. Photosynthetically active radiation (PAR) determines the photosynthetic activity of the seagrass plant that determines how much carbon dioxide is fixed, how much dissolved organic carbon (DOC) is exuded from the leaves and root system, and how much oxygen is transported into the rhizosphere. Oxygen transportation into the rhizosphere alters the redox conditions in the rhizosphere, differentiating it from the surrounding sediments that are usually anoxic and sulfidic.[75][76]

Seagrass holobiont[edit]

The concept of the holobiont, which emphasizes the importance and interactions of a microbial host with associated microorganisms and viruses and describes their functioning as a single biological unit,[77] has been investigated and discussed for many model systems, although there is substantial criticism of a concept that defines diverse host-microbe symbioses as a single biological unit.[78] The holobiont and hologenome concepts have evolved since the original definition,[79] and there is no doubt that symbiotic microorganisms are pivotal for the biology and ecology of the host by providing vitamins, energy and inorganic or organic nutrients, participating in defense mechanisms, or by driving the evolution of the host.[80] Although most work on host-microbe interactions has been focused on animal systems such as corals, sponges, or humans, there is a substantial body of literature on plant holobionts.[81] Plant-associated microbial communities impact both key components of the fitness of plants, growth and survival,[82] and are shaped by nutrient availability and plant defense mechanisms.[83] Several habitats have been described to harbor plant-associated microbes, including the rhizoplane (surface of root tissue), the rhizosphere (periphery of the roots), the endosphere (inside plant tissue), and the phyllosphere (total above-ground surface area).[75]

Seagrass meadows[edit]

Seagrass beds/meadows can be either monospecific (made up of a single species) or in mixed beds. In temperate areas, usually one or a few species dominate (like the eelgrass Zostera marina in the North Atlantic), whereas tropical beds usually are more diverse, with up to thirteen species recorded in the Philippines.

Seagrass beds are diverse and productive ecosystems, and can harbor hundreds of associated species from all phyla, for example juvenile and adult fish, epiphytic and free-living macroalgae and microalgae, mollusks, bristle worms, and nematodes. Few species were originally considered to feed directly on seagrass leaves (partly because of their low nutritional content), but scientific reviews and improved working methods have shown that seagrass herbivory is an important link in the food chain, feeding hundreds of species, including green turtles, dugongs, manatees, fish, geese, swans, sea urchins and crabs. Some fish species that visit/feed on seagrasses raise their young in adjacent mangroves or coral reefs.

Seagrasses trap sediment and slow down water movement, causing suspended sediment to settle out. Trapping sediment benefits coral by reducing sediment loads, improving photosynthesis for both coral and seagrass.[84]

Seagrass bed with several echinoids
Seagrass bed with dense turtle grass (Thalassia testudinum) and an immature queen conch (Eustrombus gigas)

Although often overlooked, seagrasses provide a number of ecosystem services.[85][86] Seagrasses are considered ecosystem engineers.[87][9][8] This means that the plants alter the ecosystem around them. This adjusting occurs in both physical and chemical forms. Many seagrass species produce an extensive underground network of roots and rhizome which stabilizes sediment and reduces coastal erosion.[88] This system also assists in oxygenating the sediment, providing a hospitable environment for sediment-dwelling organisms.[87] Seagrasses also enhance water quality by stabilizing heavy metals, pollutants, and excess nutrients.[89][9][8] The long blades of seagrasses slow the movement of water which reduces wave energy and offers further protection against coastal erosion and storm surge. Furthermore, because seagrasses are underwater plants, they produce significant amounts of oxygen which oxygenate the water column. These meadows account for more than 10% of the ocean's total carbon storage. Per hectare, it holds twice as much carbon dioxide as rain forests and can sequester about 27.4 million tons of CO2 annually.[90]

Seagrass meadows provide food for many marine herbivores. Sea turtles, manatees, parrotfish, surgeonfish, sea urchins and pinfish feed on seagrasses. Many other smaller animals feed on the epiphytes and invertebrates that live on and among seagrass blades.[91] Seagrass meadows also provide physical habitat in areas that would otherwise be bare of any vegetation. Due to this three dimensional structure in the water column, many species occupy seagrass habitats for shelter and foraging. It is estimated that 17 species of coral reef fish spend their entire juvenile life stage solely on seagrass flats.[92] These habitats also act as a nursery grounds for commercially and recreationally valued fishery species, including the gag grouper (Mycteroperca microlepis), red drum, common snook, and many others.[93][94] Some fish species utilize seagrass meadows and various stages of the life cycle. In a recent publication, Dr. Ross Boucek and colleagues discovered that two highly sought after flats fish, the common snook and spotted sea trout provide essential foraging habitat during reproduction.[95] Sexual reproduction is extremely energetically expensive to be completed with stored energy; therefore, they require seagrass meadows in close proximity to complete reproduction.[95] Furthermore, many commercially important invertebrates also reside in seagrass habitats including bay scallops (Argopecten irradians), horseshoe crabs, and shrimp. Charismatic fauna can also be seen visiting the seagrass habitats. These species include West Indian manatee, green sea turtles, and various species of sharks. The high diversity of marine organisms that can be found on seagrass habitats promotes them as a tourist attraction and a significant source of income for many coastal economies along the Gulf of Mexico and in the Caribbean.

Diminishing meadows[edit]

The storage of carbon is an essential ecosystem service as we move into a period of elevated atmospheric carbon levels. However, some climate change models suggest that some seagrasses will go extinct – Posidonia oceanica is expected to go extinct, or nearly so, by 2050.[citation needed]

The UNESCO world heritage site around the Balearic islands of Mallorca and Formentera includes about 55,000 hectares (140,000 acres) of Posidonia oceanica, which has global significance because of the amount of carbon dioxide it absorbs. However, the meadows are being threatened by rising temperatures, which slows down its growth, as well as damage from anchors.[96]

See also[edit]

References[edit]

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Further references[edit]

  • den Hartog, C. 1970. The Sea-grasses of the World. Verhandl. der Koninklijke Nederlandse Akademie van Wetenschappen, Afd. Natuurkunde, No. 59(1).
  • Duarte, Carlos M. and Carina L. Chiscano “Seagrass biomass and production: a reassessment” Aquatic Botany Volume 65, Issues 1–4, November 1999, Pages 159–174.
  • Green, E.P. & Short, F.T.(eds). 2003. World Atlas of Seagrasses. University of California Press, Berkeley, CA. 298 pp.
  • Hemminga, M.A. & Duarte, C. 2000. Seagrass Ecology. Cambridge University Press, Cambridge. 298 pp.
  • Hogarth, Peter The Biology of Mangroves and Seagrasses (Oxford University Press, 2007)
  • Larkum, Anthony W.D., Robert J. Orth, and Carlos M. Duarte (Editors) Seagrasses: Biology, Ecology and Conservation (Springer, 2006)
  • Orth, Robert J. et al. "A Global Crisis for Seagrass Ecosystems" BioScience December 2006 / Vol. 56 No. 12, Pages 987–996.
  • Short, F.T. & Coles, R.G.(eds). 2001. Global Seagrass Research Methods. Elsevier Science, Amsterdam. 473 pp.
  • A.W.D. Larkum, R.J. Orth, and C.M. Duarte (eds). Seagrass Biology: A Treatise. CRC Press, Boca Raton, FL, in press.
  • A. Schwartz; M. Morrison; I. Hawes; J. Halliday. 2006. Physical and biological characteristics of a rare marine habitat: sub-tidal seagrass beds of offshore islands. Science for Conservation 269. 39 pp. [1]
  • Waycott, M, McMahon, K, & Lavery, P 2014, A guide to southern temperate seagrasses, CSIRO Publishing, Melbourne

External links[edit]