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Seaweed fertiliser is
organic fertilizer Organic may refer to: * Organic, of or relating to an organism, a living entity * Organic, of or relating to an anatomical organ Chemistry * Organic matter, matter that has come from a once-living organism, is capable of decay or is the product ...
made from
seaweed Seaweed, or macroalgae, refers to thousands of species of macroscopic, multicellular, marine algae. The term includes some types of ''Rhodophyta'' (red), '' Phaeophyta'' (brown) and ''Chlorophyta'' (green) macroalgae. Seaweed species such as ...
that is used in agriculture to increase soil fertility and plant growth. The use of seaweed fertilizer dates back to antiquity and has a broad array of benefits for the soils. Seaweed fertilizer can be applied in a number of different forms, including refined liquid
extract An extract (essence) is a substance made by extracting a part of a raw material, often by using a solvent such as ethanol, oil or water. Extracts may be sold as tinctures or absolutes or dried and powdered. The aromatic principles of ma ...
s and dried, pulverized organic material. Through its composition of various bioactive molecules, seaweed functions as a strong
soil conditioner A soil conditioner is a product which is added to soil to improve the soil’s physical qualities, usually its fertility (ability to provide nutrition for plants) and sometimes its mechanics. In general usage, the term "soil conditioner" is often ...
, bio-remediator, and
biological pest control Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or ot ...
, with each seaweed
phylum In biology, a phylum (; : phyla) is a level of classification, or taxonomic rank, that is below Kingdom (biology), kingdom and above Class (biology), class. Traditionally, in botany the term division (taxonomy), division has been used instead ...
offering various benefits to soil and crop health. These benefits can include increased tolerance to abiotic stressors, improved soil texture and water retention, and reduced occurrence of diseases. On a broader socio-ecological scale, seaweed
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. Nelu ...
and fertilizer development have significant roles in biogeochemical
nutrient cycling A nutrient cycle (or ecological recycling) is the movement and exchange of inorganic and organic matter back into the production of matter. Energy flow is a unidirectional and noncyclic pathway, whereas the movement of mineral nutrients is cyc ...
through carbon storage and the uptake of nitrogen and phosphorus. Seaweed fertilizer application to soils can also alter the structure and function of
microbial A microorganism, or microbe, is an organism of microscopic size, which may exist in its single-celled form or as a colony of cells. The possible existence of unseen microbial life was suspected from antiquity, with an early attestation in ...
communities. Seaweed aquaculture has the potential to yield
ecosystem service Ecosystem services are the various benefits that humans derive from ecosystems. The interconnected living and non-living components of the natural environment offer benefits such as pollination of crops, clean air and water, decomposition of wast ...
s by providing a source of nutrition to human communities and a mechanism for improving water quality in natural systems and aquaculture operations. The rising popularity of organic farming practices is drawing increased attention towards the various applications of seaweed-derived fertilizers and soil additives. While the seaweed fertilizer industry is still in its infancy, it holds significant potential for sustainable economic development as well as the reduction of nutrient runoff in coastal systems. There are however ongoing challenges associated with the use and production of seaweed fertilizer including the spread of diseases and
invasive species An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...
, the risk of heavy-metal accumulation, and the efficiency and refinement of production methods.


Nomenclature and taxonomy

“Seaweed" is one of the common names given to multicellular macroalgae, such as green algae (
Chlorophyta Chlorophyta is a division of green algae informally called chlorophytes. Description Chlorophytes are eukaryotic organisms composed of cells with a variety of coverings or walls, and usually a single green chloroplast in each cell. They are ...
), brown algae (
Phaeophyceae Brown algae (: alga) are a large group of multicellular algae comprising the class Phaeophyceae. They include many seaweeds located in colder waters of the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and polar regio ...
), and red algae (
Rhodophyta Red algae, or Rhodophyta (, ; ), make up one of the oldest groups of eukaryotic algae. The Rhodophyta comprises one of the largest phyla of algae, containing over 7,000 recognized species within over 900 genera amidst ongoing taxonomic revisions. ...
). The term, seaweed is sometimes used to refer to
microalgae Microalgae or microphytes are microscopic scale, microscopic algae invisible to the naked eye. They are phytoplankton typically found in freshwater and marine life, marine systems, living in both the water column and sediment. They are unicellul ...
and plants as well. Seaweeds are typically
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
organisms which have a structure called a holdfast, that keeps them anchored to the sea floor; they also have a stipe, otherwise known as a stem, and blade-shaped foliage. Sargassum seaweed is one exception to this anatomy and function, as it does not attach to the benthic environment. The color of seaweeds generally follows depth/light, with green seaweeds, brown seaweeds, and red seaweeds corresponding to shallow, moderate, and deeper waters respectively; red seaweeds are sometimes found up to 30 meters in depth. The smallest seaweeds grow only a few millimeters in height, while the largest seaweeds can grow up to 50 meters in height. There are an estimated 1,800 green, 1,800 brown, and 6,200 red seaweed species in existence. Brown seaweeds are generally known as kelp, but are also known by other common names such as rockweed and wracks. Red seaweeds are the most diverse group of seaweed, and along with green seaweeds, are most closely related to terrestrial plants, whereas brown seaweeds are the most distantly related to terrestrial plants. Seaweeds are found extensively in shallow natural environments, and farmed both in the ocean and in land-based
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. Nelu ...
operations. Most brown seaweeds that are found in the wild are from the genera ''
Laminaria ''Laminaria'' is a genus of brown algae, brown seaweed in the order Kelp, Laminariales (kelp), comprising 31 species native to the north Atlantic and northern Pacific Oceans. This economically important genus is characterized by long, leathery L ...
'', '' Undaria'', '' Hizikia,'' whereas most brown seaweeds that are farmed for uses such as fertilizer and heavy metal indication, are from the species ''
Ascophyllum ''Ascophyllum nodosum'' is a large, common cold water seaweed or Phaeophyceae, brown alga (Phaeophyceae) in the Family (biology), family Fucaceae. Its common names include knotted wrack, egg wrack, feamainn bhuí, rockweed, knotted kelp and Norweg ...
'', ''
Ecklonia ''Ecklonia'' is a genus of kelp (brown algae) belonging to the family Lessoniaceae. The genus name of ''Ecklonia'' is in honour of Christian Friedrich Ecklon (1795–1868), who was a Danish botanical collector and apothecary. The genus was cir ...
'', ''
Fucus ''Fucus'' is a genus of brown algae found in the intertidal zones of rocky seashores almost throughout the world. Description and life cycle The thallus is perennial with an irregular or disc-shaped holdfast or with haptera. The erect portion ...
'', ''
Sargassum ''Sargassum'' is a genus of brown macroalgae ( seaweed) in the order Fucales of the Phaeophyceae class. Numerous species are distributed throughout the temperate and tropical oceans of the world, where they generally inhabit shallow water and ...
''. Green seaweeds that are used as bioindicators, for heavy metal indication for example, are from the genera ''Ulva'' and ''Enteromorpha''. Red seaweed from the genus ''Poryphora,'' is commonly used for human food.


History

The first written record of agricultural use seaweed was from ancient Greek and Roman civilizations in the 2nd century, where foraged beach castings were used to feed livestock and wrap plant roots for preservation. However,
stable isotope analysis Isotope analysis is the identification of isotopic signature, abundance of certain stable isotopes of chemical elements within organic and inorganic compounds. Isotopic analysis can be used to understand the flow of energy through a food web ...
of prehistoric sheep teeth in
Orkney Orkney (), also known as the Orkney Islands, is an archipelago off the north coast of mainland Scotland. The plural name the Orkneys is also sometimes used, but locals now consider it outdated. Part of the Northern Isles along with Shetland, ...
indicate that early peoples used seaweed as livestock
fodder Fodder (), also called provender (), is any agriculture, agricultural foodstuff used specifically to feed domesticated livestock, such as cattle, domestic rabbit, rabbits, sheep, horses, chickens and pigs. "Fodder" refers particularly to food ...
over 5,000 years ago, and researchers speculate that foraged seaweed was also used as fertilizer because ashed remnants of seaweed were found in archeological sites. Such agricultural techniques might have been key to the survival of early settlements in Scotland. Historical records and archaeological evidence of seaweed fertilizer use in the coastal
Atlantic The Atlantic Ocean is the second largest of the world's five oceanic divisions, with an area of about . It covers approximately 17% of Earth's surface and about 24% of its water surface area. During the Age of Discovery, it was known for se ...
are vast and scattered, ranging from
Scandinavia Scandinavia is a subregion#Europe, subregion of northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. ''Scandinavia'' most commonly refers to Denmark, Norway, and Sweden. It can sometimes also ...
to
Portugal Portugal, officially the Portuguese Republic, is a country on the Iberian Peninsula in Southwestern Europe. Featuring Cabo da Roca, the westernmost point in continental Europe, Portugal borders Spain to its north and east, with which it share ...
, from the
Neolithic The Neolithic or New Stone Age (from Ancient Greek, Greek 'new' and 'stone') is an archaeological period, the final division of the Stone Age in Mesopotamia, Asia, Europe and Africa (c. 10,000 BCE to c. 2,000 BCE). It saw the Neolithic Revo ...
period through the 20th century. Most details of seaweed fertilizer use come from the
British Isles The British Isles are an archipelago in the Atlantic Ocean, North Atlantic Ocean off the north-western coast of continental Europe, consisting of the islands of Great Britain, Ireland, the Isle of Man, the Inner Hebrides, Inner and Outer Hebr ...
,
Channel Islands The Channel Islands are an archipelago in the English Channel, off the French coast of Normandy. They are divided into two Crown Dependencies: the Jersey, Bailiwick of Jersey, which is the largest of the islands; and the Bailiwick of Guernsey, ...
, Normandy and
Brittany Brittany ( ) is a peninsula, historical country and cultural area in the north-west of modern France, covering the western part of what was known as Armorica in Roman Gaul. It became an Kingdom of Brittany, independent kingdom and then a Duch ...
(France), where a variety of application techniques were used over the centuries, and some continue to this day. Ireland has a long history (12th century) of harvesting seaweed for fertilizing nutrient-poor post glacial soils using composted
manure Manure is organic matter that is used as organic fertilizer in agriculture. Most manure consists of animal feces; other sources include compost and green manure. Manures contribute to the fertility of soil by adding organic matter and nut ...
as enrichment and the increased agricultural productivity allowed the Irish population to grow substantially. The Channel Islands (12th century) used a dried blend of red and brown seaweeds, called "Vraic" or " wrack", to spread over potato fields during the winter months to enrich before planting the crop in the spring. Similarly, coastal people in Normandy and Brittany have been collecting "wrack" using wood rakes since the neolithic period, though the fertilizer composition originally included all
marine debris Marine debris, also known as marine litter, is human-created solid material that has deliberately or accidentally been released in seas or the ocean. Floating oceanic debris tends to accumulate at the center of gyres and on coastlines, freque ...
that washed ashore. In 17th–19th century
Scotland Scotland is a Countries of the United Kingdom, country that is part of the United Kingdom. It contains nearly one-third of the United Kingdom's land area, consisting of the northern part of the island of Great Britain and more than 790 adjac ...
, ''Fucus spp''. were cultivated by placing rocky substrate in the
intertidal The intertidal zone or foreshore is the area above water level at low tide and underwater at high tide; in other words, it is the part of the littoral zone within the tidal range. This area can include several types of habitats with various sp ...
zones to encourage seaweed settlement. The seaweed biomass was then used in composted trenches, where crops (potatoes, oats, wheat, onions) were grown directly in the sandy fertilizer mixture. This ‘
lazy bed Lazy bed ( or ; ; Faroese language, Faroese: ''letivelta'') is a traditional method of arable cultivation, often used for potatoes. Rather like cord rig cultivation, parallel banks of ridge and furrow are dug by spade although lazy beds have ...
’ method afforded minimal crop rotation and allowed rugged landscape and acidic soils to be farmed, where plant growth was otherwise unsuitable. The high value of seaweed in these regions caused political disputes over harvesting rights and in Ireland such rights were established before the country itself. These early applications of seaweed fertilizer were limited to coastlines, where the macroalgae could be harvested from the intertidal or collected after a storm washed it to shore. However, dried wrack mixtures or ashed ‘fucus’
potash Potash ( ) includes various mined and manufactured salts that contain potassium in water- soluble form.
could be transported further inland because it weighs less than wet seaweed. Seaweed fertilizer spread inland when a
kelp Kelps are large brown algae or seaweeds that make up the order (biology), order Laminariales. There are about 30 different genus, genera. Despite its appearance and use of photosynthesis in chloroplasts, kelp is technically not a plant but a str ...
industry developed in Scotland, Norway, and Brittany in the 18th and 19th century. The industry developed out of demand for ashed soda, or potash, which was used to create glass and soap, and led to shortages for agricultural applications in traditional coastal communities. Potash is a water-soluble
potassium Potassium is a chemical element; it has Symbol (chemistry), symbol K (from Neo-Latin ) and atomic number19. It is a silvery white metal that is soft enough to easily cut with a knife. Potassium metal reacts rapidly with atmospheric oxygen to ...
rich concentrate made from plant matter, so it was also exported as a fertilizer. Coastal communities in the seaweed industry both expanded and struggled to keep up with the demand. Early commercial kelp export in Scotland devastated traditional agriculture in the region because intensive labor was needed during the seaweed growing season to harvest and process the kelp, which led to a labor transition from farming to kelp processing. Additionally, exploitation of kelp resources for potash production left little kelp behind for local fertilizer and coastal land became more desirable than inland regions. The Scottish seaweed industry went through multiple boom and bust cycles, employing 10,000 families and producing 3,000 tonnes of ash per year during its peak. The export price of kelp ash dropped in 1822, leading to a sudden emigration from the area because the crop was no longer profitable enough to support such a large industry. Kelp exploitation and toxic ash processing caused ecological and economic damage in Orkney and left many people sick and blinded. The kelp industry picked up again for
iodine Iodine is a chemical element; it has symbol I and atomic number 53. The heaviest of the stable halogens, it exists at standard conditions as a semi-lustrous, non-metallic solid that melts to form a deep violet liquid at , and boils to a vi ...
production in 1845, and
alginate Alginic acid, also called algin, is a naturally occurring, edible polysaccharide found in brown algae. It is hydrophilic and forms a viscous gum when hydrated. When the alginic acid binds with sodium and calcium ions, the resulting salts are k ...
(a thickening agent) production in the early 1900s, which reinvigorated kelp harvest. Global production of seaweed fertilizer largely phased out when chemical fertilizers were developed in the 1920s, due to the cheaper production cost. Chemical fertilizers revolutionized the agriculture industry and allowed the human population to grow far beyond the limits of traditional food production methods. Synthetic fertilizers are still the predominant global source for commercial agricultural applications due to the cheap cost of production and widespread access. However, small scale organic farmers and coastal communities continued traditional seaweed techniques in regions with a rich seaweed history. The first industrial kelp liquid fertilizer, Maxicrop, was created by Reginald Milton in 1947. The creation of liquid fertilizer has allowed for more widespread application of seaweed-derived fertilizer to inland regions and sparked a growing
agronomic Agricultural economics is an applied field of economics concerned with the application of economic theory in optimizing the production and distribution of food and fiber products. Agricultural economics began as a branch of economics that specif ...
interest in seaweed for a variety of agricultural applications, including foliage spray, biostimulants, and soil conditioning. Interestingly, the historic rise of seaweed
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. Nelu ...
did not align with fertilizer production because the European countries that produce seaweed fertilizer haven't developed a significant aquaculture industry;
seaweed farming Seaweed farming or kelp farming is the practice of aquaculture, cultivating and harvesting seaweed. In its simplest form farmers gather from natural beds, while at the other extreme farmers fully control the crop's biological life cycle, life c ...
is also currently dominated by China and Indonesia, where the crop is grown for food and other lucrative uses.


Aquaculture

The development of modern seaweed
mariculture Mariculture, sometimes called marine farming or marine aquaculture, is a branch of aquaculture involving the cultivation of marine organisms for food and other animal products, in seawater. Subsets of it include ( offshore mariculture), fish fa ...
/aquaculture has allowed the expansion of seaweed fertilizer research and improved processing methods since the 1950s. Seaweed has been cultivated in Asian countries for food production for centuries, but seaweed aquaculture is now growing rapidly across the world for specialty use in
biofuel Biofuel is a fuel that is produced over a short time span from Biomass (energy), biomass, rather than by the very slow natural processes involved in the formation of fossil fuels such as oil. Biofuel can be produced from plants or from agricu ...
,
agar Agar ( or ), or agar-agar, is a jelly-like substance consisting of polysaccharides obtained from the cell walls of some species of red algae, primarily from " ogonori" and " tengusa". As found in nature, agar is a mixture of two components, t ...
, cosmetics, medicine, and
bioplastic Bioplastics are plastic materials produced from renewable biomass sources. Timeline of plastic development, Historically, bioplastics made from natural materials like shellac or Celluloid, cellulose had been the first plastics. Since the end of ...
s. The nascent agricultural seaweed sector, including animal feed, soil additives, and
agrochemical An agrochemical or agrichemical, a contraction of ''agricultural chemical'', is a chemical product used in industrial agriculture. Agrichemical typically refers to biocides (pesticides including insecticides, herbicides, fungicides and nematicide ...
s, makes up less than 1% of the overall global value of seaweed aquaculture. However, significant interest in agricultural applications of the crop has increased dramatically since 1950, as specialty agrochemical uses for seaweed materials have been demonstrated through scientific research. Increased concern over the depletion and degradation of marine resources in the past century, coupled with the threats of
climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
, has increased global interest in sustainable solutions for blue economic development of the oceans. Seaweed aquaculture is promoted as a solution to expand novel industry development and food security while simultaneously restoring damaged ecosystems. Unlike terrestrial crops, growing seaweed requires no land, feed, fertilizers, pesticides, and water resources. Different seaweeds also offer a variety of
ecosystem service Ecosystem services are the various benefits that humans derive from ecosystems. The interconnected living and non-living components of the natural environment offer benefits such as pollination of crops, clean air and water, decomposition of wast ...
s (discussed below), which contribute to the growing popularity of seaweed as a
bioremediation Bioremediation broadly refers to any process wherein a biological system (typically bacteria, microalgae, fungi in mycoremediation, and plants in phytoremediation), living or dead, is employed for removing environmental pollutants from air, wate ...
crop. Fertilizer plays an important role in sustainable seaweed aquaculture development because seaweed farming can help alleviate excess nutrient loading associated with terrestrial chemical fertilizer run-off and applying organic seaweed fertilizer on soil closes the nutrient loop between land and sea. Additionally, seaweed fertilizer can be produced using by-products from other industries or raw materials that are unsuitable for human consumption, such as rotting or infected biomass or biowaste products from carrageenan processing methods. Seaweed aquaculture is also important for supporting sustainble growth of the seaweed fertilizer industry because it limits the potential for exploitation of native seaweed for commercial interests. However, the nascent seaweed aquaculture industry faces a number of challenges to sustainable development, as discussed below. Environmental impacts of seaweed harvest and production need to be carefully scrutinized to protect coastal communities and maintain the socioeconomic benefits of using seaweed resources in industry.


Ecosystem services

Seaweed mariculture for purposes including fertilizer production, has the potential to improve environmental conditions in coastal habitats, especially with regards to toxic algal blooms, as mariculture seaweeds uptake excess nutrients that have resulted from runoff, thereby inhibiting the growth of toxic algal blooms that harm local ecosystems. Seaweed fertilizers can also be more biodegradable, less toxic, and less hazardous than chemical fertilizers, depending on the type of seaweed fertilizer. Seaweeds are used in aquaculture operations to uptake fish waste as nutrients and improve water quality parameters. Humans use seaweeds nutritionally as food, industrially for animal feed and plant fertilizer, and ecologically to improve environmental conditions. Seaweeds have been consumed by humans for centuries because they have excellent nutritional profiles, contain minerals,
trace element __NOTOC__ A trace element is a chemical element of a minute quantity, a trace amount, especially used in referring to a micronutrient, but is also used to refer to minor elements in the composition of a rock, or other chemical substance. In nutr ...
s,
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...
s, and
vitamin Vitamins are Organic compound, organic molecules (or a set of closely related molecules called vitamer, vitamers) that are essential to an organism in small quantities for proper metabolism, metabolic function. Nutrient#Essential nutrients, ...
s, and are high in fiber and low in calories. Red seaweeds have the highest protein content and brown seaweeds have the lowest protein content. Of all the red seaweeds, ''Porphyra'', is the genus most frequently used for human consumption. Brown seaweeds are so plentiful that they most used for industrial animal feeds and fertilizers. Furthermore, seaweeds are currently being investigated as a potential source of sustainable biofuel, as well as being investigated as a potential component of wastewater treatment, because some species are able to absorb and remove heavy metals and other toxicants from water bodies, and also generally serve as water quality indicators.


Ecosystem impacts

Any ecosystem impacts of using seaweed for plant and crop fertilizer are primarily due to how the seaweed is harvested. Large-scale, unsustainable seaweed farming can lead to the displacement and alteration of native habitats due to the presence of farming infrastructure in the water, and day-to-day anthropogenic operations in the area. Seaweed is currently harvested from farmed sources, wild sources, and from beach collection efforts. Harvesting wild seaweed will tend to have negative impacts on local ecosystems, especially if existing populations are overexploited and rendered unable to provide
ecosystem services Ecosystem services are the various benefits that humans derive from Ecosystem, ecosystems. The interconnected Biotic_material, living and Abiotic, non-living components of the natural environment offer benefits such as pollination of crops, clean ...
. There is also a risk that large, industrial scale seaweed monocultures will be established in natural benthic environments, leading to the competitive exclusion of native seaweeds and sea grasses, which inhabit the depths underneath seaweed farms. Furthermore, large, industrial scale seaweed farming can alter the natural benthic environment that they are established in, by altering environmental parameters such as light availability, the movement of water, sedimentation rates and nutrient levels, and due to the general, overall stress caused by anthropogenic factors.


Production and application methods

Brown seaweeds are most commonly used for fertilizer production, at present and historically. Seaweed fertilizer can be used as a crude addition to soil as
mulch A mulch is a layer of material applied to the surface of soil. Reasons for applying mulch include conservation of soil moisture, improving soil fertility, fertility and health of the soil, reducing Weed control, weed growth, and enhancing the v ...
, composted to break down the hardy raw material, or dried and pulverized to make the nutrients more bioavailable to plant roots. Compost fertilization is a technique that any small-scale organic farm can readily use if they have access to seaweed, though extracts are more common for large-scale commercial applications. Commercial manufacturing processes are often more technical than traditional techniques using raw biomass and use different biochemical processes to concentrate and extract the most beneficial nutrients from seaweed. A simple liquid fertilizer can be created by
fermenting Fermentation is a type of anaerobic metabolism which harnesses the redox potential of the reactants to make adenosine triphosphate (ATP) and organic end products. Organic molecules, such as glucose or other sugars, are catabolized and reduced ...
seaweed leaves in water, though the process is intensified and hastened industrially through heat and pressure. Other methods for liquid extraction include a soft-extraction with low temperature milling to suspend fine particles in water, heating the raw material with alkaline sodium or potassium to extract nutrients and the addition of enzymes to aid in biochemical decomposition. Extraction of bioavailable nutrients from raw seaweed is achieved by breaking down the hardy
cell wall A cell wall is a structural layer that surrounds some Cell type, cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, ...
s through physical techniques, such as
ultrasound Ultrasound is sound with frequency, frequencies greater than 20 Hertz, kilohertz. This frequency is the approximate upper audible hearing range, limit of human hearing in healthy young adults. The physical principles of acoustic waves apply ...
extraction, boiling, or freeze-thaw. Biological fermentation techniques are also used to degrade the cells. Physical extraction techniques are often faster, but more expensive and result in poorer crop yield in trials. Since seaweed extract has
chelating Chelation () is a type of bonding of ions and their molecules to metal ions. It involves the formation or presence of two or more separate coordinate bonds between a polydentate (multiple bonded) ligand and a single central metal atom. These l ...
properties that maintain trace metal ions bioavailability to plants, additional
micronutrient Micronutrients are essential chemicals required by organisms in small quantities to perform various biogeochemical processes and regulate physiological functions of cells and organs. By enabling these processes, micronutrients support the heal ...
s are often added to solution to increase the fertilization benefit to specific crops. Organic fertilization techniques have lower environmental consequences in comparison to the production of artificial chemical fertilizers, because they use no harsh caustic or organic
solvent A solvent (from the Latin language, Latin ''wikt:solvo#Latin, solvō'', "loosen, untie, solve") is a substance that dissolves a solute, resulting in a Solution (chemistry), solution. A solvent is usually a liquid but can also be a solid, a gas ...
s to produce fertilizer and the seaweed raw material is a renewable resource, as opposed to mineral deposits and fossil fuels needed to synthesize chemical fertilizer. Large-scale agricultural use of synthetic fertilizer depletes soil fertility and increases water hardness over time, so recent trends in agricultural development are following an organic approach to sustain food production through improved soil management and bio-fertilization techniques. Seaweed extracts are bio-fertilizers that can also be used as biostimulants, which are applied to enhance nutrient efficiency and abiotic stress tolerance. New extraction technologies are being developed to improve efficiency and target the isolation of specific compounds for specialized applications of seaweed biostimulants, though specific extraction techniques are frequently trade secrets. Additionally, many liquid fertilizer extraction processes can complement other industrial uses for seaweed, such as
carrageenan Carrageenans or carrageenins ( ; ) are a family of natural linear sulfation, sulfated polysaccharides. They are extracted from red algae, red edible seaweeds. Carrageenans are widely used in the food industry, for their gelling, thickening, an ...
production, which increases the economic benefit of the same seaweed crop.


Nutrient cycling

To support a growing seaweed aquaculture industry many studies have evaluated the nutrient cycle dynamics of different seaweed species in addition to exploring co-production applications including bioremediation and carbon sequestration. Seaweeds can form highly productive communities in coastal regions, dominating the nutrient cycles within these ecosystems. As primary producers, seaweeds incorporate inorganic carbon, light, and nutrients (such as nitrogen and phosphorus), into biomass through photosynthesis. Harvesting seaweed from marine environments results in the net removal of these elements from these ecosystems in addition to the removal of heavy metals and contaminants. For photosynthesis, seaweeds utilize both inorganic nitrogen, in the forms of
nitrate Nitrate is a polyatomic ion with the chemical formula . salt (chemistry), Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are solubility, soluble in wa ...
- NO3,
ammonium Ammonium is a modified form of ammonia that has an extra hydrogen atom. It is a positively charged (cationic) polyatomic ion, molecular ion with the chemical formula or . It is formed by the protonation, addition of a proton (a hydrogen nucleu ...
- NH4+, and organic nitrogen in the form of
urea Urea, also called carbamide (because it is a diamide of carbonic acid), is an organic compound with chemical formula . This amide has two Amine, amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest am ...
. Primary production using nitrate is generally considered new production because nitrate is externally supplied through
upwelling Upwelling is an physical oceanography, oceanographic phenomenon that involves wind-driven motion of dense, cooler, and usually nutrient-rich water from deep water towards the ocean surface. It replaces the warmer and usually nutrient-depleted sur ...
and riverine input, and often has been converted from forms of nitrogen that are released from biological respiration. However, primary production using ammonium is denoted as recycled production because ammonium is internally supplied through regeneration by
heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
s within ecosystems. For example, the ammonium excreted by fish and invertebrates within the same coastal ecosystems as seaweeds can support seaweed production through providing a nitrogen source. Phosphorus is supplied inorganically as
phosphate Phosphates are the naturally occurring form of the element phosphorus. In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthop ...
(PO43-) and generally follows similar seasonal patterns to nitrate. In addition, seaweeds require inorganic carbon, typically supplied from the environment in the form of carbon dioxide (CO2) or
bicarbonate In inorganic chemistry, bicarbonate (IUPAC-recommended nomenclature: hydrogencarbonate) is an intermediate form in the deprotonation of carbonic acid. It is a polyatomic anion with the chemical formula . Bicarbonate serves a crucial bioche ...
(HCO3). Similar to other marine photosynthesizing organisms like
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater Aquatic ecosystem, ecosystems. The name comes from the Greek language, Greek words (), meaning 'plant', and (), mea ...
, seaweeds also experience nutrient limitations impacting their ability to grow. Nitrogen is the most commonly found limiting nutrient for seaweed photosynthesis, although phosphorus has also been found to be limiting. The ratio of inorganic carbon, nitrogen, and phosphorus is also important to ensure balanced growth. Generally the N:P ratio for seaweeds is 30:1, however, the ratio can differ significantly among species and requires experimental testing to identify the specific ratio for a given species. Exploring the relationship between nutrient cycling and seaweed growth is vital to optimizing seaweed aquaculture and understanding the functions and benefits of seaweed applications, including its use as a fertilizer, bio-remediator, and in the blue economy.


Coastal eutrophication

A growing population and intensification of industry and agriculture have increased the volume of wastewater discharged into coastal marine ecosystems. These waters typically contain high concentrations of nitrogen and phosphorus, and relatively high heavy metal concentrations, leading to
eutrophication Eutrophication is a general term describing a process in which nutrients accumulate in a body of water, resulting in an increased growth of organisms that may deplete the oxygen in the water; ie. the process of too many plants growing on the s ...
of many coastal ecosystems. Eutrophication results from the excessive nutrient load within these ecosystems resulting from the pollution of waters entering the oceans from industry, animal feed, and synthetic fertilizers, and thus over-fertilizes these systems. Eutrophication leads to high productivity in coastal systems, which can result in coastal hypoxia and
ocean acidification Ocean acidification is the ongoing decrease in the pH of the Earth's ocean. Between 1950 and 2020, the average pH of the ocean surface fell from approximately 8.15 to 8.05. Carbon dioxide emissions from human activities are the primary cause of ...
, two major concerns for coastal ecosystems. A notable service of seaweed farming is its ability to act as a bio-remediator through uptake and removal of th excessive nutrients in the coastal ecosystems with their application to land uses. Brown algae, due in part to their large size, have been noted for their high productivity and corresponding high nutrient uptake in coastal ecosystems. Additionally, studies have focused on how brown algae growth can be optimized to increase biomass production and therefore increase the quantity of nutrients removed from these ecosystems. Studies have also explored the potential of brown algae to sequester large volumes of carbon (
blue carbon Blue carbon is a concept within climate change mitigation that refers to "biologically driven carbon fluxes and storage in marine systems that are amenable to management". Most commonly, it refers to the role that tidal marshes, mangroves and Seag ...
).


Bio-remediation in eutrophic ecosystems

Seaweeds have received significant attention for their potential to mitigate eutrophication in coastal ecosystems through nutrient uptake during primary production in
integrated multi-trophic aquaculture Integrated multi-trophic aquaculture (IMTA) is a type of aquaculture where the By-product, byproducts, including waste, from one aquatic species are used as inputs (fertilizers, food) for another. Farmers combine fed aquaculture (e.g., fish, shri ...
(IMTA). Bioremediation involves the use of biological organisms to lower the concentrations of nitrogen, phosphorus, and heavy metal concentrations in marine ecosystems. The bioremediation potential of seaweeds depends, in part, on their growth rate which is controlled by numerous factors including water movement, light,
desiccation Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. The ...
, temperature, salinity, life stage, and age class. It has also been proposed that in eutrophic ecosystems phosphorus can become limiting to seaweed growth due to the high N:P ratio of the wastewater entering these ecosystems. Bioremediation practices have been widely used due to their cost-effective ability to reduce excess nutrients in coastal ecosystems leading to a decrease in
harmful algal bloom A harmful algal bloom (HAB), or excessive algae growth, sometimes called a red tide in marine environments, is an algal bloom that causes negative impacts to other organisms by production of natural algae-produced toxins, water deoxygenation, ...
s and an oxygenation of the water column. Seaweeds have also been studied for their potential use in the
biosorption Biosorption is a physiochemical process that occurs naturally in certain biomass which allows it to passively concentrate and bind contaminants onto its cellular structure. Biosorption can be defined as the ability of biological materials to accumu ...
and accumulation of heavy metals in polluted waters, although the accumulation of heavy metals may impact algal growth.


Blue carbon

Blue carbon methods involve the use of marine ecosystems for carbon storage and burial. Seaweed aquaculture shows potential to act as a CO2 sink through the uptake of carbon during photosynthesis, transformation of inorganic carbon into biomass, and ultimately the fixation of carbon which can later be exported and buried. Duarte et al. (2017) outline a potential strategy for a seaweed farming blue carbon initiative. However the contribution of seaweed to blue carbon has faced controversy over the ability of seaweed to act as a net sink for atmospheric carbon. Krause-Jensen et al., (2018) discuss two main criteria for seaweed farming to be considered a blue carbon initiative: it must be both extensive in size and sequestration rate and possess the ability to be actionable by humans, that the sequestration rate can be managed by human action. Seaweed farming, including the use of seaweed as fertilizer could become an important contributor in climate mitigation strategies through carbon sequestration and storage.


Functions and benefits of seaweed fertilizer


Fertilization

Seaweed functions as an organic bio-fertilizer. Because seaweed is rich in micro and
macronutrients A nutrient is a substance used by an organism to survive, grow and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi and protists. Nutrients can be incorporated into cells for metabolic purposes or excret ...
, humic acids, and
phytohormones Plant hormones (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, including embryogenesis, the regulation of organ size ...
, it enhances
soil fertility Soil fertility refers to the ability of soil to sustain agricultural plant growth, i.e. to provide plant habitat and result in sustained and consistent yields of high quality.
. In addition, seaweed-derived fertilizers contain
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long-chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wat ...
s,
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
s, and
fatty acid In chemistry, in particular in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated and unsaturated compounds#Organic chemistry, saturated or unsaturated. Most naturally occurring fatty acids have an ...
s which improve the moisture and nutrient retention of soil, contributing to improved crop growth. More trace minerals are found in seaweed than those produced with animal byproducts. The application of seaweed fertilizers can also result in enhanced tolerance to abiotic stressors that generally inhibit crop growth and yield such as low moisture, high salinity, and freezing temperatures. These stress tolerance benefits appear to be driven by physiological changes induced in crops by the seaweed, including improved energy storage, enhanced root morphology, and greater metabolic potential, enhancing the plant's ability to survive unfavorable conditions. '' Kappaphycus alvarezzi'' extracts have also resulted in considerable reductions in the leakage of
electrolyte An electrolyte is a substance that conducts electricity through the movement of ions, but not through the movement of electrons. This includes most soluble Salt (chemistry), salts, acids, and Base (chemistry), bases, dissolved in a polar solven ...
s, as well as enhanced
chlorophyll Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...
and
carotenoid Carotenoids () are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, archaea, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, corn, tomatoes, cana ...
production, and water content. Research has also demonstrated that wheat plants treated with seaweed extracts have accumulated key
osmoprotectant Osmoprotectants or compatible solutes are small organic molecules with neutral charge and low toxicity at high concentrations that act as osmolytes and help organisms to survive in extreme osmotic stress. Osmoprotectants can be placed in three che ...
s such as
proline Proline (symbol Pro or P) is an organic acid classed as a proteinogenic amino acid (used in the biosynthesis of proteins), although it does not contain the amino group but is rather a secondary amine. The secondary amine nitrogen is in the p ...
, other amino acids, and total protein. Foliar applications of seaweed fertilizer extract have been shown to improve the uptake of nitrogen, phosphorus, potassium, and sulfur in soybeans such as G''lycine max''. Research has also demonstrated that brown algae seaweed extracts can improve tomato plant growth, overall crop yield, and resistance to environmental stressors. Additional documented benefits of using seaweed as a fertilizer include reduced transplant shock, increased leaf surface area, and increased sugar content.


Soil conditioning

As a soil conditioner, seaweed fertilizer can improve the physical qualities of soil, such as aeration and water retention. Clay soils that lack organic matter and
porosity Porosity or void fraction is a measure of the void (i.e. "empty") spaces in a material, and is a fraction of the volume of voids over the total volume, between 0 and 1, or as a percentage between 0% and 100%. Strictly speaking, some tests measure ...
benefit from the
humic acid Humic substances (HS) are colored relatively recalcitrant organic compounds naturally formed during long-term decomposition and transformation of biomass residues. The color of humic substances varies from bright yellow to light or dark brown lead ...
and soluble alginates found in seaweed. These compounds bond with metallic radicals which cause the
clay Clay is a type of fine-grained natural soil material containing clay minerals (hydrous aluminium phyllosilicates, e.g. kaolinite, ). Most pure clay minerals are white or light-coloured, but natural clays show a variety of colours from impuriti ...
particles to aggregate, thereby improving the texture, aeration, and retention of the soil by stimulating clay disaggregation. The degradation of alginates also supplements the soil with organic matter, enhancing its fertility. In particular, brown seaweeds such as ''Sargassum'' are known to have valuable soil conditioning properties. This seaweed contains soluble alginates as well as alginic acid, which catalyzes the bacterial decomposition of organic matter. This process improves
soil quality Soil quality refers to the condition of soil based on its capacity to perform ecosystem services that meet the needs of human and non-human life.Tóth, G., Stolbovoy, V. and Montanarella, 2007. Soil Quality and Sustainability Evaluation - An integ ...
by enhancing populations of
nitrogen-fixing Nitrogen fixation is a chemical process by which molecular dinitrogen () is converted into ammonia (). It occurs both biologically and abiological nitrogen fixation, abiologically in chemical industry, chemical industries. Biological nitrogen ...
bacteria and by supplementing the soil with additional conditioners through the waste products produced by these bacteria.


Bio-remediation of polluted soils

Seaweed functions as a bio-remediator through its
adsorption Adsorption is the adhesion of atoms, ions or molecules from a gas, liquid or dissolved solid to a surface. This process creates a film of the ''adsorbate'' on the surface of the ''adsorbent''. This process differs from absorption, in which a ...
of harmful pollutants.
Functional group In organic chemistry, a functional group is any substituent or moiety (chemistry), moiety in a molecule that causes the molecule's characteristic chemical reactions. The same functional group will undergo the same or similar chemical reactions r ...
s on the algal surface such as
ester In chemistry, an ester is a compound derived from an acid (either organic or inorganic) in which the hydrogen atom (H) of at least one acidic hydroxyl group () of that acid is replaced by an organyl group (R). These compounds contain a distin ...
, hydroxyl, carbonyl amino, sulfhydryl, and phosphate groups drive the biosorption of heavy metal ions. Seaweeds such as ''Gracilaria corticata varcartecala'' and ''Grateloupia lithophila'' effectively remove a wide variety of heavy metals, including
chromium Chromium is a chemical element; it has Symbol (chemistry), symbol Cr and atomic number 24. It is the first element in Group 6 element, group 6. It is a steely-grey, Luster (mineralogy), lustrous, hard, and brittle transition metal. Chromium ...
(III) and (IV), mercury (II), lead (II), and cadmium (II) from their environment. In addition, ''Ulva spp.'' and '' Gelidium spp.'' have been shown to enhance the degradation of
DDT Dichlorodiphenyltrichloroethane, commonly known as DDT, is a colorless, tasteless, and almost odorless crystalline chemical compound, an organochloride. Originally developed as an insecticide, it became infamous for its environmental impacts. ...
in polluted soils and may reduce its bioavailability. Although there is significant potential for seaweed to serve as a bio-remediator for polluted soils, more research required to fully develop the mechanisms for this process in the context of the agriculture. Heavy metals accumulated by seaweed fertilizer may transfer to crops in some cases, causing significant implications for the public health. The application of
biochar Biochar is a form of charcoal, sometimes modified, that is intended for organic use, as in soil. It is the lightweight black remnants remaining after the pyrolysis of biomass, consisting of carbon and ashes. Despite its name, biochar is steril ...
is another strategy that can remediate and enhance infertile soils. Seaweed can be transformed into biochar and used as a means of increasing the organic matter and nutrient content of the soil. Different types of seaweed appear to yield unique nutrients and parameters; red seaweeds, for example, create
biochar Biochar is a form of charcoal, sometimes modified, that is intended for organic use, as in soil. It is the lightweight black remnants remaining after the pyrolysis of biomass, consisting of carbon and ashes. Despite its name, biochar is steril ...
that is rich in
potassium Potassium is a chemical element; it has Symbol (chemistry), symbol K (from Neo-Latin ) and atomic number19. It is a silvery white metal that is soft enough to easily cut with a knife. Potassium metal reacts rapidly with atmospheric oxygen to ...
and
sulfur Sulfur ( American spelling and the preferred IUPAC name) or sulphur ( Commonwealth spelling) is a chemical element; it has symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms ...
and is more acidic than biochar generated from brown seaweeds. While this is a new field of research, current data shows that targeted breeding of seaweeds may result in
biochar Biochar is a form of charcoal, sometimes modified, that is intended for organic use, as in soil. It is the lightweight black remnants remaining after the pyrolysis of biomass, consisting of carbon and ashes. Despite its name, biochar is steril ...
s that can be tailored to different types of soil and crops.


Integrated pest management

The addition of seaweed to soil can increase crop health and resistance to diseases. Seaweeds contain a diverse array of bioactive molecules that can respond to diseases and pests, including
steroid A steroid is an organic compound with four fused compound, fused rings (designated A, B, C, and D) arranged in a specific molecular configuration. Steroids have two principal biological functions: as important components of cell membranes t ...
s,
terpene Terpenes () are a class of natural products consisting of compounds with the formula (C5H8)n for n ≥ 2. Terpenes are major biosynthetic building blocks. Comprising more than 30,000 compounds, these unsaturated hydrocarbons are produced predomi ...
s,
acetogenin Acetogenins are a class of polyketide natural products found in plants of the family Annonaceae. They are characterized by linear 32- or 34-carbon chains containing oxygenated functional groups including hydroxyls, ketones, epoxides, tetrahyd ...
s, and amino acid-derived
polymer A polymer () is a chemical substance, substance or material that consists of very large molecules, or macromolecules, that are constituted by many repeat unit, repeating subunits derived from one or more species of monomers. Due to their br ...
s. The application of seaweed extracts reduces the presence of harmful pests including
nematode The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...
s and insects. While the application of seaweed seems to reduce the harmful effects of nematode infestation, the combination of seaweed application and
carbofuran Carbofuran is a carbamate insecticide, banned in the US, the EU and Canada but still widely used in South America, Australia and Asia. It is a systemic insecticide, which means that the plant absorbs it through the roots, and from there the pla ...
, a chemical
nematocide A nematicide is a type of chemical pesticide used to kill plant-parasitic nematodes. Nematicides have tended to be broad-spectrum toxicants possessing high volatility or other properties promoting migration through the soil. Aldicarb (Temik), a ...
, seems to be most effective. In addition, several species of seaweed appear to hinder the early growth and development of numerous detrimental insects, including ''Sargassum swartzii'', '' Padina pavonica'', and ''Caulerpa denticulata''.


Soil microbial response to seaweed fertilizer treatment

Shifts in bacterial and fungal communities, in response to seaweed fertilizer treatment, have only recently been studied. Soil Microbial community composition and functionality is largely driven by underlying soil health and abiotic properties. Many DNA sequencing and
omics Omics is the collective characterization and quantification of entire sets of biological molecules and the investigation of how they translate into the structure, function, and dynamics of an organism or group of organisms. The branches of scien ...
-based approaches, combined with
greenhouse A greenhouse is a structure that is designed to regulate the temperature and humidity of the environment inside. There are different types of greenhouses, but they all have large areas covered with transparent materials that let sunlight pass an ...
experiments, have been used to characterize microbial responses to seaweed fertilizer treatment on a wide variety of crops. Deep
16S ribosomal RNA 16S ribosomal RNA (or 16 S rRNA) is the RNA component of the 30S subunit of a prokaryotic ribosome ( SSU rRNA). It binds to the Shine-Dalgarno sequence and provides most of the SSU structure. The genes coding for it are referred to as 16S ...
(rRNA)
amplicon In molecular biology, an amplicon is a piece of DNA or RNA that is the source and/or product of amplification or replication events. It can be formed artificially, using various methods including polymerase chain reactions (PCR) or ligase chain ...
sequencing of the bacteria found in the soils of tomato plots, treated with a ''
Sargassum horneri ''Sargassum horneri'' is a species of brown macroalgae that is common along the coast of Japan and Korea. It is an annual algae which has a varying fertile season along the coast. In Wakasa Bay, Japan it begins to grow in early autumn through wi ...
'' fermented seaweed fertilizer, showed a large shift in
alpha diversity In ecology, alpha diversity (α-diversity) is the mean species diversity in a site at a local scale. The term was introduced by R. H. WhittakerWhittaker, R. H. (1960) Vegetation of the Siskiyou Mountains, Oregon and California. Ecological Monogra ...
and
beta diversity In ecology, beta diversity (β-diversity or true beta diversity) is the ratio between regional and local species diversity. The term was introduced by Robert Whittaker (ecologist), R. H. Whittaker together with the terms alpha diversity (α-diversi ...
indices between untreated soils and soils after 60 days. This shift in community composition was correlated with a 1.48-1.83 times increase in tomato yield in treated soils. Though dominant bacterial phyla remained similar between treatment groups, changes in the abundance of the class,
Bacilli Bacilli is a Taxonomy (biology), taxonomic Class (biology), class of bacteria that includes two orders, Bacillales and Lactobacillales, which contain several well-known pathogens such as ''Bacillus anthracis'' (the cause of anthrax). ''Bacilli'' ...
and family, ''
Micrococcaceae ''Micrococcaceae'', from Ancient Greek μικρός (''mikrós''), meaning "small", and κόκκος (''kókkos''), meaning "sphere", includes bacterial genera of Gram positive cocci that inhabit the air and skin, such as '' Micrococcus luteus ...
'' were noted. Enzyme assays also displayed an increase in
protease A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalysis, catalyzes proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the formation of new protein products ...
,
polyphenol oxidase Polyphenol oxidase (PPO; also polyphenol oxidase i, chloroplastic), an enzyme involved in fruit browning, is a tetramer that contains four atoms of copper per molecule. PPO may accept monophenols and/or ''o''-diphenols as substrates. The ...
,
dehydrogenase A dehydrogenase is an enzyme belonging to the group of oxidoreductases that oxidizes a substrate by reducing an electron acceptor, usually NAD+/NADP+ or a flavin coenzyme such as FAD or FMN. Like all catalysts, they catalyze reverse as well as ...
,
invertase β-Fructofuranosidase is an enzyme that catalyzes the hydrolysis (breakdown) of the table sugar sucrose into fructose and glucose. Sucrose is a fructoside. Alternative names for β-fructofuranosidase include invertase, saccharase, glucosucrase ...
, and
urease Ureases (), functionally, belong to the superfamily of amidohydrolases and phosphotriesterases. Ureases are found in numerous Bacteria, Archaea, fungi, algae, plants, and some invertebrates. Ureases are nickel-containing metalloenzymes of high ...
activity, which was thought to be induced by microbial community alterations. Each of the microbial and enzymatic results listed above were noted to improve the nutrient turnover and quality in soils treated with fertilizer. To investigate interactions between plant growth-promoting rhizobacteria (PGPR) and seaweed-derived extract, Ngoroyemoto et al. treated '' Amaranthus hybridus'' with both Kelpak and PGPR and measured impacts on plant growth. It was found that the treatment of plants with Kelpak® and the bacteria, ''
Pseudomonas fluorescens ''Pseudomonas fluorescens'' is a common Gram-negative, rod-shaped bacterium. It belongs to the ''Pseudomonas'' genus; 16S rRNA analysis as well as phylogenomic analysis has placed ''P. fluorescens'' in the ''P. fluorescens'' group within the genu ...
'' and ''
Bacillus licheniformis ''Bacillus licheniformis'' is a bacterium commonly found in the soil. It is found on bird feathers, especially chest and back plumage, and most often in ground-dwelling birds (like sparrows) and aquatic species (like ducks). It is a gram-positi ...
'', decreased plant stress responses and increased production. The most recently mentioned study provides implications for crop benefits when the application of seaweed fertilizer to soils favors the growth of PGPR. Wang et al. found that apple
seedling A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle (embryonic root), the hypocotyl (embry ...
s treated with seaweed fertilizer differed markedly in fungal diversity and species richness, when compared to no-treatment control groups. These findings were complemented by increases in soil quality and enzyme activities in treated soil groups, which supports the hypothesis that the fertilizer promoted the growth of plant-beneficial fungal species. With the use of 16S rRNA and fungal
internal transcribed spacer Internal transcribed spacer (ITS) is the spacer DNA situated between the small-subunit ribosomal RNA (rRNA) and large-subunit rRNA genes in the chromosome or the corresponding transcribed region in the polycistronic rRNA precursor transcript. ...
(ITS) sequencing, Renaut et al. examined the effect of ''
Ascophyllum nodosum ''Ascophyllum nodosum'' is a large, common cold water seaweed or brown alga ( Phaeophyceae) in the family Fucaceae. Its common names include knotted wrack, egg wrack, feamainn bhuí, rockweed, knotted kelp and Norwegian kelp. It grows only in the ...
'' extract treatment on the
rhizosphere The rhizosphere is the narrow region of soil or Substrate (biology), substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Pore space in soil, Soil pores in the rhizosphere can ...
s of pepper and tomato plants in greenhouses. This group found that bacterial and fungal
species composition Relative species abundance is a component of biodiversity and is a measure of how common or rare a species is relative to other species in a defined location or community.Hubbell, S. P. 2001. ''The unified neutral theory of biodiversity and biogeog ...
and community structures differed based on treatment. A rise of the abundance of certain
amplicon sequence variant An amplicon sequence variant (ASV) is any one of the inferred single DNA sequences recovered from a high-throughput analysis of marker genes. Because these analyses, also called "amplicon reads," are created following the removal of erroneous sequ ...
s (ASVs) were also directly correlated with increases in plant health and growth. These ASVs included fungi in the family, '' Microascaceae'', the genus, '' Mortierella spp.'', and several other uncultured ASVs. A large diversity of bacterial ASVs were identified to be positively correlated with growth in this same study, including ''
Rhizobium ''Rhizobium'' is a genus of Gram-negative soil bacteria that fix nitrogen. ''Rhizobium'' species form an endosymbiotic nitrogen-fixing association with roots of (primarily) legumes and other flowering plants. The bacteria colonize plant ce ...
'', ''
Sphingomonas ''Sphingomonas'' was defined in 1990 as a group of Gram-negative, rod-shaped, chemoheterotrophic, strictly Aerobic organism, aerobic bacteria. They possess ubiquinone 10 as their major Electron transport chain#Quinone carriers, respiratory quinon ...
'', '' Sphingobium'', and ''
Bradyrhizobium ''Bradyrhizobium'' is a genus of Gram-negative bacteria, Gram-negative soil bacteria, many of which fix nitrogen. Nitrogen fixation is an important part of the nitrogen cycle. Plants cannot use atmospheric nitrogen (N2); they must use nitrogen co ...
''.


Resistance to plant pathogens

Application of the seaweed fertilizer may also increase resistance to plant pathogens. In greenhouse samples, Ali et al. tested the treatment of ''
Ascophyllum nodosum ''Ascophyllum nodosum'' is a large, common cold water seaweed or brown alga ( Phaeophyceae) in the family Fucaceae. Its common names include knotted wrack, egg wrack, feamainn bhuí, rockweed, knotted kelp and Norwegian kelp. It grows only in the ...
'' extract on tomato and sweet pepper crops and found that it both increased plant health and reduced the incidence of plant pathogens. Further investigation showed that the up-regulation of pathogen defense-related enzymes led to the reduction of the pathogens, '' Xanthomonas campestris pv. vesicatoria'' and '' Alternaria solani''. Chen et al. found that ''Ascophyllum nodosum'' treatment positively impacted the community composition of maize rhizospheres. This may have critical implications for plant health because the structure of rhizosphere microbial communities can aid in the resistance of plants to soil-borne pathogens. Other pathogen reductions include the mitigation of carrot foliar fungal diseases following ''
Ascophyllum nodosum ''Ascophyllum nodosum'' is a large, common cold water seaweed or brown alga ( Phaeophyceae) in the family Fucaceae. Its common names include knotted wrack, egg wrack, feamainn bhuí, rockweed, knotted kelp and Norwegian kelp. It grows only in the ...
'' treatment and inoculation with the fungal pathogens, '' Alternaria radicina'' and ''
Botrytis cinerea ''Botrytis cinerea'' is a necrotrophic fungus that affects many plant species, although its most notable hosts may be wine grapes. In viticulture, it is commonly known as "botrytis bunch rot"; in horticulture, it is usually called "grey mould" ...
''. Reduced disease severity was noted at 10 and 20 days post-inoculation in comparison to control plants, and the seaweed treatment was found to be more effective at reducing disease pathology than
salicylic acid Salicylic acid is an organic compound with the formula HOC6H4COOH. A colorless (or white), bitter-tasting solid, it is a precursor to and a active metabolite, metabolite of acetylsalicylic acid (aspirin). It is a plant hormone, and has been lis ...
, a known plant protector from biotic and abiotic stresses. Islam et al. had similar results when treating ''
Arabidopsis thaliana ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small plant from the mustard family (Brassicaceae), native to Eurasia and Africa. Commonly found along the shoulders of roads and in disturbed land, it is generally ...
'' with brown algal extracts, followed by inoculation with the fungal pathogen ''
Phytophthora cinnamomi ''Phytophthora cinnamomi'', also known as cinnamon fungus, is a soil-borne water mould that produces an infection which causes a condition in plants variously called "dieback", "root rot", or (in certain '' Castanea'' species), "ink disease". O ...
''. This group analyzed plant RNA transcripts and found that the seaweed extract primed ''A. thaliana'' to defend against the fungal pathogen before its inoculation, which led to increased host survival and decreased susceptibility to infection. Fewer studies have analyzed the impact of seaweed fertilizer treatment on plant resistance to viral pathogens, however limited auspicious results have been demonstrated. It has been shown that green, brown, and red seaweeds contain
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long-chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wat ...
s that elicit
pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
response pathways in plants, which primes defense against viruses, along with bacteria and fungi. Specifically, defense enzymes, including
phenylalanine ammonia lyase The enzyme phenylalanine ammonia lyase (EC 4.3.1.24) catalyzes the conversion of L-phenylalanine to ammonia and ''trans''-cinnamic acid.: :L-phenylalanine = ''trans''-cinnamate + NH3 Phenylalanine ammonia lyase (PAL) is the first and committe ...
and
lipoxygenase Lipoxygenases () (LOX) are a family of (non- heme) iron-containing enzymes, more specifically oxidative enzymes, most of which catalyze the dioxygenation of polyunsaturated fatty acids in lipids containing a cis,cis-1,4- pentadiene into ce ...
, are activated and lead to viral defense. Aqueous and ethanolic extracts from the brown alga, ''
Durvillaea antarctica ''Durvillaea antarctica'', also known as ' and ', is a large, robust species of southern bull kelp found on the coasts of Chile, southern New Zealand, and Macquarie Island.Smith, J.M.B. and Bayliss-Smith, T.P. (1998). Kelp-plucking: coastal eros ...
'' was shown to decrease pathological symptoms of
tobacco mosaic virus Tobacco mosaic virus (TMV) is a positive-sense single-stranded RNA virus species in the genus '' Tobamovirus'' that infects a wide range of plants, especially tobacco and other members of the family Solanaceae. The infection causes characteris ...
(TMV) in tobacco leaves. Another study done on tobacco plants found that sulfated fucan
oligosaccharide An oligosaccharide (; ) is a carbohydrate, saccharide polymer containing a small number (typically three to ten) of monosaccharides (simple sugars). Oligosaccharides can have many functions including Cell–cell recognition, cell recognition and ce ...
s, extracted from brown algae, induced local and
systemic acquired resistance Systemic acquired resistance (SAR) is a "whole-plant" resistance response that occurs following an earlier localized exposure to a pathogen. SAR is analogous to the innate immune system found in animals, and although there are many shared aspects b ...
to TMV. Based on the above results, it can be stated that the application of seaweed fertilizers has considerable potential to provide broad benefits to the agricultural crops and resistance to bacterial, fungal, and viral plant
pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
s.


References

{{Reflist Wikipedia Student Program Organic fertilizers Seaweeds