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Extracellular phototropic digestion is a process in which
saprobionts Saprobionts are organisms that digest their food externally and then absorb the products. This process is called saprotrophic nutrition. Fungi are examples of saprobiontic organisms, which are a type of decomposer. Saprobiontic organisms feed off ...
feed by secreting
enzymes Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different molecule ...
through the
cell membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment ( ...
onto the food. The enzymes catalyze the digestion of the food ie diffusion, transport,
osmotrophy Osmotrophy is a feeding mechanism involving the movement of dissolved organic compounds by osmosis for nutrition. Organisms that use osmotrophy are called osmotrophs. Some mixotrophic microorganisms use osmotrophy to derive some of their energy. ...
or
phagocytosis Phagocytosis () is the process by which a cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs phagocytosis is ...
. Since
digestion Digestion is the breakdown of large insoluble food molecules into small water-soluble food molecules so that they can be absorbed into the watery blood plasma. In certain organisms, these smaller substances are absorbed through the small intest ...
occurs outside the cell, it is said to be extracellular. It takes place either in the lumen of the
digestive system The human digestive system consists of the gastrointestinal tract plus the accessory organs of digestion (the tongue, salivary glands, pancreas, liver, and gallbladder). Digestion involves the breakdown of food into smaller and smaller compone ...
, in a gastric cavity or other digestive organ, or completely outside the body. During extracellular digestion, food is broken down outside the cell either mechanically or with acid by special molecules called enzymes. Then the newly broken down nutrients can be absorbed by the cells nearby. Humans use extracellular digestion when they eat. Their teeth grind the food up, enzymes and acid in the
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
liquefy it, and additional enzymes in the
small intestine The small intestine or small bowel is an organ in the gastrointestinal tract where most of the absorption of nutrients from food takes place. It lies between the stomach and large intestine, and receives bile and pancreatic juice through the p ...
break the food down into parts their cells can use. Extracellular digestion is a form of digestion found in all saprobiontic
annelids The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecolo ...
,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s,
lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.chordates A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These five ...
, including
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
.


In fungi

Fungi are
heterotrophic 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 ...
organisms. Heterotrophic nutrition means that fungi utilize extracellular sources of organic energy,
organic material Organic matter, organic material, or natural organic matter refers to the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have c ...
or organic matter, for their maintenance, growth and reproduction. Energy is derived from the breakdown of the chemical bond between carbon and either carbon or other components of compounds such as a
phosphate ion 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 orthophosphoric acid . The phosphate or orthophosphate ion is derived from phospho ...
. The extracellular sources of energy may be
simple sugar Monosaccharides (from Greek ''monos'': single, '' sacchar'': sugar), also called simple sugars, are the simplest forms of sugar and the most basic units (monomers) from which all carbohydrates are built. They are usually colorless, water-solub ...
s,
polypeptide Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides. A p ...
s or more
complex carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or may ...
. Fungi can only absorb small molecules through their walls. For fungi to gain their energy needs, they find and absorb organic molecules appropriate to their needs, either immediately or following some form of enzyme diminution outside the
thallus Thallus (plural: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. Many of these organisms wer ...
. The small molecules are then absorbed, used directly or reconstituted (transformed) into organic molecules within the cell. When a skeletonized leaf is seen in the litter, it is because
recalcitrant Recalcitrant seeds are seeds that do not survive drying and freezing during ex-situ conservation. By and large, these seeds cannot resist the effects of drying or temperatures less than 10 °C (50 °F); thus, they cannot be stored for long periods ...
materials remain and digestion is continuing. The fungi that utilize a variety of energy sources usually absorb the simplest compounds first, then the more complex. For instance, the formation of
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
is repressed by high concentrations of
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, using ...
in the
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...
. On depletion of primary sources of glucose, enzymes to degrade more complex molecules such as cellulose and
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
, are then released. Thus soluble sugars and
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...
s are removed first from a leaf released from a tree. Starch is then broken down and absorbed. Subsequently,
pectin Pectin ( grc, πηκτικός ': "congealed" and "curdled") is a heteropolysaccharide, a structural acid contained in the primary lamella, in the middle lamella, and in the cell walls of terrestrial plants. The principal, chemical component of ...
and cellulose are digested. Finally,
wax Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to giv ...
es are degraded and
lignin Lignin is a class of complex organic polymers that form key structural materials in the support tissues of most plants. Lignins are particularly important in the formation of cell walls, especially in wood and bark, because they lend rigidity ...
oxidized Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or a d ...
. The staggering of energy acquisition results in the efficient utilization of available energy.


Detection of digestive enzymes in fungi

The regulation of nutrient acquisition appears to be controlled by general phenomena. Only a small group of enzymes, mostly
hydrolase Hydrolase is a class of enzyme that commonly perform as biochemical catalysts that use water to break a chemical bond, which typically results in dividing a larger molecule into smaller molecules. Some common examples of hydrolase enzymes are este ...
s, can be detected in the culture filtrate of well-fed fungi. This suggests that specific inducers control the manufacture and release of enzymes for degradation. The most common complex carbohydrate available in the environment is cellulose. In the absence of glucose, detection of cellulose, for instance, induces the expression of celluloses. As a consequence, fungi specifically target the breakdown of the cellulose in their environment, and do not waste energy on the unnecessary formation of enzymes for degradation of molecules that may not be present. Fungi have an efficient process to gain energy. Because of the huge range of potential food sources, fungi have evolved enzymes suitable for the environments in which they are usually found. The range of enzymes, though wide in many species, is not sufficient for survival in all environments. Fungi require other competitive attributes to ensure continued survival. The opposite is also true. Some fungi have highly specific metabolic capabilities which enable occupation of specific habitats, utilizing molecules which are unavailable to other fungi. Further, utilization of a common and abundant substrate has led many fungi to evolve a range of highly specific degradative enzymes. Among the fungi are species that are generalist in their nutrient requirements, some that have specific nutrient requirements, and many that are in between.


Excretion of digestive enzymes

Enzymes are manufactured close to the hyphal tip. Some are packaged in vesicles associated with the Golgi and then delivered to the hyphal tip. The contents are released at the tip. Some enzymes are actively excreted through the
plasma membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
, where they diffuse through or act in the cell wall. Note that the enzymes released from the hyphal tip require an aqueous environment for release and subsequent degradative activity.


Absorption of digested products

The molecules absorbed through the plasma membrane tend to be smaller than 5,000 Da, so only simple sugars, amino acids,
fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, fr ...
s and other small molecules can be taken up following digestion. The molecules are taken up in solution. In some cases, the molecules are processed by enzymes located within the cell wall. For instance, sucrose inverters have been localized in walls of yeasts. Glucose appears to be the sugar preferred by most fungi. Uptake of other sugars is repressed when glucose is available. Similarly,
ammonium The ammonium cation is a positively-charged polyatomic ion with the chemical formula or . It is formed by the protonation of ammonia (). Ammonium is also a general name for positively charged or protonated substituted amines and quaternary a ...
,
glutamine Glutamine (symbol Gln or Q) is an α-amino acid that is used in the biosynthesis of proteins. Its side chain is similar to that of glutamic acid, except the carboxylic acid group is replaced by an amide. It is classified as a charge-neutral, ...
and
asparagine Asparagine (symbol Asn or N) is an α-amino acid that is used in the biosynthesis of proteins. It contains an α-amino group (which is in the protonated −NH form under biological conditions), an α-carboxylic acid group (which is in the depro ...
regulate the uptake of nitrogen compounds, and
cysteine Cysteine (symbol Cys or C; ) is a semiessential proteinogenic amino acid with the formula . The thiol side chain in cysteine often participates in enzymatic reactions as a nucleophile. When present as a deprotonated catalytic residue, sometime ...
of sulphur compounds.


Joint intracellular and extracellular digestion in cnidarians

Joint intracellular and extracellular digestion In hydra and other cnidarians, the food is caught by the tentacles and ingested through the mouth into the single large digestive cavity, the
gastrovascular cavity The gastrovascular cavity is the primary organ of digestion and circulation in two major animal phyla: the Coelenterates or cnidarians (including jellyfish and corals) and Platyhelminthes (flatworms). The cavity may be extensively branched into ...
. Enzymes are secreted from the cells bordering this cavity and poured on the food for extracellular digestion. Small particles of the partially digested food are engulfed into the
vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
s of the digestive cells for
intracellular digestion Every organism requires energy to be active.Anderson, O. Roger (1 January 1970). "Intracellular Digestion". ''The American Biology Teacher''. 32 (8): 461–467. doi:10.2307/4443206. However, to obtain energy from its outside environment, cells m ...
. Any undigested and un-absorbed food is finally thrown out of the mouth.


Invert digestive systems are bags and tubes

Single-celled organism A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms an ...
s as well as
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...
s digest their food intracellularly. Other multi-cellular organisms digest their food extracellularly, within a digestive cavity. In this case the
digestive enzyme Digestive enzymes are a group of enzymes that break down polymeric macromolecules into their smaller building blocks, in order to facilitate their absorption into the cells of the body. Digestive enzymes are found in the digestive tracts of anim ...
s are released into a cavity that is continuous with the animal's external environment. In cnidarians and in
flatworm The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegment ...
s such as
planarian A planarian is one of the many flatworms of the traditional class Turbellaria. It usually describes free-living flatworms of the order Tricladida (triclads), although this common name is also used for a wide number of free-living platyhelmint ...
s, the digestive cavity, called a gastrovascular cavity, has only one opening that serves as both mouth and anus. There is no specialization within this type of digestive system because every cell is exposed to all stages of food digestion. Specializing occurs when the digestive tract or alimentary canal has a separate mouth and anus so that transport of food is one-way. The most primitive digestive tract is seen in
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s (phylum Nematode), where it is simply a tubular gut lined by an
epithelial Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
membrane. Earthworms (phylum Annelids) have a digestive tract specialized in different regions for the ingestion, storage, fragmentation, digestion and absorption of food. All more complex animal groups, including all vertebrates, show similar specializations. The ingested food may be stored in a specialized region of the digestive tract or subjected to physical fragmentation. This fragmentation may occur through the chewing action of teeth (in the mouth of many vertebrates) or the grinding action of pebbles (in the gizzard of earthworms and birds). Chemical digestion then occurs, breaking down the larger food molecules of
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 wa ...
s and
disaccharide A disaccharide (also called a double sugar or ''biose'') is the sugar formed when two monosaccharides are joined by glycosidic linkage. Like monosaccharides, disaccharides are simple sugars soluble in water. Three common examples are sucrose, lact ...
s, fats, and proteins into their smallest sub-units. Chemical digestion involves
hydrolysis Hydrolysis (; ) is any chemical reaction in which a molecule of water breaks one or more chemical bonds. The term is used broadly for substitution reaction, substitution, elimination reaction, elimination, and solvation reactions in which water ...
reactions that liberate the sub unit molecules—primarily
monosaccharide Monosaccharides (from Greek ''monos'': single, '' sacchar'': sugar), also called simple sugars, are the simplest forms of sugar and the most basic units (monomers) from which all carbohydrates are built. They are usually colorless, water-solub ...
s, amino acids and fatty acids—from the food. These products of chemical digestion pass through the epithelial lining of the gut into the blood, in a process known as absorption. Any molecules in the food that are not absorbed cannot be used by the animal. These waste products are excreted, or defecated from the anus.


Extracellular digestion in other animals


Annelids

The echiuran gut is long and highly convoluted, and there is no gut in pogonophoran adults. Among other annelids, the gut is linear and unsegmented, with a mouth opening on the
peristomium The peristomium is the first true body segment in an annelid worm's body in the anterior end. It is directly behind the prostomium and contains the mouth, tentacular cirri, and sometimes feeding palps, which may instead occur on the prostomium. If ...
and an anus opening at the posterior end of the animal (
pygidium The pygidium (plural pygidia) is the posterior body part or shield of crustaceans and some other arthropods, such as insects and the extinct trilobites. In groups other than insects, it contains the anus and, in females, the ovipositor. It is compo ...
). Food is moved through the gut by cilia and/or by
muscular contraction Muscle contraction is the activation of tension-generating sites within muscle cells. In physiology, muscle contraction does not necessarily mean muscle shortening because muscle tension can be produced without changes in muscle length, such as ...
s. Digestion is primarily extracellular, although some species show an intracellular component as well.


Arthropods

The
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
digestive system is divisible into three areas: the fore gut, mid gut, and hind gut. All free-living species exhibit a distinct and separate mouth and anus, and in all species, food must be moved through the digestive tract by muscular activity rather than
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
activity since the lumen of the fore gut and hind gut is lined with
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
. Digestion is generally extracellular. Nutrients are distributed to the tissues through the hemal system.


Molluscs

Most
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
s have a complete digestive system with a separate mouth and anus. The mouth leads into a short
esophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
which leads to a stomach. Associated with the stomach are one or more
digestive gland The hepatopancreas, digestive gland or midgut gland is an organ of the digestive tract of arthropods and molluscs. It provides the functions which in mammals are provided separately by the liver and pancreas, including the production of digestive e ...
s or digestive caeca. Digestive enzymes are secreted into the lumen of these glands. Additional extracellular digestion takes place in the stomach. In
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s, digestion is entirely extracellular. In the most other mollusks, the terminal stages of digestion are completed intracellularly, within the tissue of the digestive glands. The absorbed nutrients enter the
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
for distribution throughout the body or are stored in the digestive glands for later use. Undigested waste pass through an intestine and out through the anus. Other aspects of food collection and processing have already been discussed where appropriate for each group.


Humans

The initial components of the
gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organ (biology), organs of the digestive syste ...
are the mouth and the
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struc ...
, which is the common passage of the oral and nasal cavities. The pharynx leads to the
esophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
, a muscular tube that delivers food to the
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
, where some preliminary digestion occurs; here, the digestion is ''extracellular''. From the stomach, food passes to the
small intestine The small intestine or small bowel is an organ in the gastrointestinal tract where most of the absorption of nutrients from food takes place. It lies between the stomach and large intestine, and receives bile and pancreatic juice through the p ...
, where a battery of digestive enzymes continue the digestive process. The products of digestion are absorbed across the wall of the intestine into the
bloodstream The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
. What remains is emptied into the
large intestine The large intestine, also known as the large bowel, is the last part of the gastrointestinal tract and of the digestive system in tetrapods. Water is absorbed here and the remaining waste material is stored in the rectum as feces before being r ...
, where some of the remaining water and minerals are absorbed; here the digestion is intracellular.


See also

*
Saprotrophic nutrition Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi (f ...


References

{{DEFAULTSORT:Extra-Cellular Digestion Eating behaviors