Extracellular phototropic digestion is a process in which
saprobionts feed by secreting
enzymes 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 (the ...
onto the food. The enzymes catalyze the digestion of the food ie diffusion, transport,
osmotrophy 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 i ...
. 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, 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 liquefy it, and additional enzymes in the
small intestine break the food down into parts their cells can use.
Extracellular digestion is a form of digestion found in all saprobiontic
annelids,
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 gro ...
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,
lichens and
chordates, including
vertebrates.
In fungi
Fungi are
heterotrophic 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. The extracellular sources of energy may be
simple sugars,
polypeptides or more
complex carbohydrate.
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. 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 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, usi ...
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. ...
. On depletion of primary sources of glucose, enzymes to degrade more complex molecules such as cellulose and
starch, 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 ...
s are removed first from a leaf released from a tree. Starch is then broken down and absorbed. Subsequently,
pectin and cellulose are digested. Finally,
waxes 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. 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 es ...
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
Substrate may refer to:
Physical layers
*Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached
** Substrate (locomotion), the surface over which an organism lo ...
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, 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, ...
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,
glutamine and
asparagine 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, s ...
of sulphur compounds.
Joint intracellular and extracellular digestion in cnidarians
Joint intracellular and extracellular digestion In hydra and other
cnidarian
Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in Fresh water, freshwater and Marine habitats, marine environments, predominantly the latter.
Their distinguishing feature is cnidocyt ...
s, the food is caught by the tentacles and ingested through the mouth into the single large digestive cavity, the
gastrovascular cavity. 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
vacuoles of the digestive cells for
intracellular digestion. Any undigested and un-absorbed food is finally thrown out of the mouth.
Invert digestive systems are bags and tubes
Single-celled organisms 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 throug ...
s digest their food intracellularly. Other
multi-cellular organisms digest their food extracellularly, within a digestive cavity. In this case the
digestive enzymes 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, unsegme ...
s such as
planarian
A planarian is one of the many flatworms of the traditional class (biology), 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-li ...
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
nematodes (phylum Nematode), where it is simply a tubular gut lined by an
epithelial 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 ...
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, ...
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, elimination, and solvation reactions in which water is the nucleophile.
Biological hydrolysis ...
reactions that liberate the sub unit molecules—primarily
monosaccharides, 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
The Echiura, or spoon worms, are a small group of marine animals. Once treated as a separate phylum, they are now considered to belong to Annelida. Annelids typically have their bodies divided into segments, but echiurans have secondarily los ...
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 and an anus opening at the posterior end of the animal (
pygidium). Food is moved through the gut by cilia and/or by
muscular contractions. 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 activity since the
lumen of the fore gut and hind gut is lined with
cuticle. Digestion is generally extracellular. Nutrients are distributed to the tissues through the hemal system.
Molluscs
Most
molluscs have a complete digestive system with a separate mouth and anus. The mouth leads into a short
esophagus which leads to a stomach. Associated with the stomach are one or more
digestive glands 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 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 ...
, which is the common passage of the oral and nasal cavities. The pharynx leads to the
esophagus, a muscular tube that delivers food to the
stomach, where some preliminary digestion occurs; here, the digestion is ''extracellular''.
From the stomach, food passes to the
small intestine, 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. What remains is emptied into the
large intestine, where some of the remaining water and minerals are absorbed; here the digestion is intracellular.
See also
*
Saprotrophic nutrition
References
{{DEFAULTSORT:Extra-Cellular Digestion
Eating behaviors