Probiotic

From Wikipedia, the free encyclopedia

A bottle of Yakult, a probiotic drink containing Lactobacillus paracasei.

Probiotics are live microorganisms intended to provide health benefits when consumed, generally by improving or restoring the gut flora. Probiotics are considered generally safe to consume, but may cause bacteria-host interactions and unwanted side effects in rare cases.

The original theory, similar to the modern concept, but not the term, is generally attributed to Nobel laureate Élie Metchnikoff, who postulated that yoghurt-consuming Bulgarian peasants lived longer lives because of that custom. In 1907, he wrote: "[T]he dependence of the intestinal microbes on the food makes it possible to adopt measures to modify the microbiota in our bodies[,] and to replace the harmful microbes by useful microbes."

A growing probiotics market has led to the need for stricter requirements for scientific substantiation of putative benefits conferred by microorganisms claimed to be probiotic. Although there are numerous claimed benefits marketed towards using consumer probiotic products, such as reducing gastrointestinal discomfort, improving immune health, relieving constipation, or avoiding the common cold, such claims are not supported by scientific evidence, and are prohibited as deceptive advertising in the United States by the Federal Trade Commission.

In a clinical setting however, some probiotics have been found to be useful in treating specific medical conditions, such as antibiotic-associated diarrhea in children and Clostridium difficile infection in adults. One concern is that probiotics taken by mouth can be destroyed by the acidic conditions of the stomach. As of 2010, a number of microencapsulation techniques were being developed to address this problem.

Definition

An October 2001 report by the World Health Organization (WHO) defines probiotics as live microorganisms that, "when administered in adequate amounts, confer a health benefit on the host." Following this definition, a working group convened by the Food and Agriculture Organization (FAO)/WHO in May 2002 issued the Guidelines for the Evaluation of Probiotics in Food. A consensus definition of the term probiotics, based on available information and scientific evidence, was adopted after the aforementioned joint expert consultation between the FAO of the United Nations and the WHO. This effort was accompanied by local governmental and supra-governmental regulatory bodies' requirements to better characterize health claims substantiations.

That first global effort was further developed in 2010; two expert groups of academic scientists and industry representatives made recommendations for the evaluation and validation of probiotic health claims. The same principles emerged from those two groups as were expressed in the "Guidelines" of FAO/WHO in 2002. This definition, though widely adopted, is not acceptable to the European Food Safety Authority (EFSA) because it embeds a health claim that is not measurable.

A group of scientific experts assembled in London on October 23, 2013, to discuss the scope and appropriate use of the term probiotic. That meeting was motivated by developments in the field that followed the formation of the 2001 definition, and the panel's conclusions were published in June 2014.

Probiotics in food

Live probiotic cultures are part of fermented dairy products, other fermented foods, and probiotic-fortified foods.

Some fermented products that contain lactic acid bacteria (LAB) include: vegetables such as pickled vegetables, kimchi, pao cai, and sauerkraut; soy products such as tempeh, miso, and soy sauce; and dairy products such as yogurt, kefir, and buttermilk.

Side effects

The manipulation of the gut microbiota is complex and may cause bacteria-host interactions. Though probiotics are considered safe, some have concerns about their safety in certain cases. Some people, such as those with immunodeficiency, short bowel syndrome, central venous catheters, cardiac valve disease and premature infants, may be at higher risk for adverse events. In severely ill people with inflammatory bowel disease there is a risk of the passage of viable bacteria from the gastrointestinal tract to the internal organs (bacterial translocation) as a consequence of bacteremia, which can cause adverse health consequences. Rarely, consumption of probiotics by children with lowered immune system function or who are already critically ill may result in bacteremia or fungemia (i.e., bacteria or fungi in the blood), which can lead to sepsis, a potentially fatal disease.

It has been suggested that Lactobacillus contributes to obesity in humans, but no evidence of this relationship has been found.

Global consumption

In 2015, the global retail market value for probiotics was US$41 billion, including sales of probiotic supplements, fermented milk products, and yogurt, which alone accounted for 75% of total consumption. Innovation in probiotic products in 2015 was mainly from supplements, which produced US$4 billion and was projected to grow 37% globally by 2020. Consumption of yogurt products in China has increased by 20% per year since 2014.

Regulation

The European Food Safety Authority has rejected all petitions by commercial manufacturers for health claims on probiotic products in Europe due to insufficient research, and thus inconclusive proof of effectiveness. Occurring over many years, the scientific reviews established that a cause-and-effect relationship had not been sufficiently proven in the products submitted. The European Commission placed a ban on putting the word "probiotic" on the packaging of products because such labeling misleads consumers to believe a health benefit is provided by the product when no scientific proof exists to demonstrate that health effect.

In the United States, the FDA and Federal Trade Commission have issued warning letters and imposed punishment on various manufacturers of probiotic products whose labels claim to treat a disease or condition. Food product labeling requires language approval by the Food and Drug Administration, so probiotic manufacturers have received warning letters for making disease or treatment claims. The Federal Trade Commission has taken punitive actions, including a US$21 million fine coordinated by 39 different state governments against a major probiotic manufacturer for deceptive advertising and exaggerated claims of health benefits for a yogurt and probiotic dairy drink.

Yogurt labeling

The National Yogurt Association (NYA) of the United States gives a Live & Active Cultures Seal to refrigerated yogurt products that contain 100 million cultures per gram, or frozen yogurt products that contain 10 million cultures per gram at the time of manufacture. In 2002, the US Food and Drug Administration (FDA) and World Health Organization recommended that “the minimum viable numbers of each probiotic strain at the end of the shelf-life” be reported on labeling, but most companies that give a number report the viable cell count at the date of manufacture, a number that could be much higher what exists at consumption. Because of the variability in storage conditions and time before eating, it is difficult to tell exactly how many or how much active culture remains at the time of consumption.

History

Probiotics have received renewed attention in the 21st century from product manufacturers, research studies, and consumers. The history of probiotics can be traced to the first use of cheese and fermented products, that were well known to the Greeks and Romans who recommended their consumption. The fermentation of dairy foods represents one of the oldest techniques for food preservation.

Élie Metchnikoff first suggested the possibility of colonizing the gut with beneficial bacteria in the early 20th century.

The original modern hypothesis of the positive role played by certain bacteria was first introduced by Russian scientist and Nobel laureate Élie Metchnikoff, who in 1907 suggested that it would be possible to modify the gut microbiota and to replace harmful microbes with useful microbes. Metchnikoff, at that time a professor at the Pasteur Institute in Paris, proposed the hypothesis that the aging process results from the activity of putrefactive (proteolytic) microbes producing toxic substances in the large bowel. Proteolytic bacteria such as clostridia, which are part of the normal gut microbiota, produce toxic substances including phenols, indols, and ammonia from the digestion of proteins. According to Metchnikoff, these compounds were responsible for what he called intestinal autointoxication, which would cause the physical changes associated with old age.

It was at that time known that milk fermented with lactic-acid bacteria inhibits the growth of proteolytic bacteria because of the low pH produced by the fermentation of lactose. Metchnikoff had also observed that certain rural populations in Europe, for example in Bulgaria and the Russian steppes, who lived largely on milk fermented by lactic-acid bacteria were exceptionally long lived. Based on these observations, Metchnikoff proposed that consumption of fermented milk would "seed" the intestine with harmless lactic-acid bacteria and decrease the intestinal pH, and that this would suppress the growth of proteolytic bacteria. Metchnikoff himself introduced in his diet sour milk fermented with the bacteria he called "Bulgarian Bacillus" and believed his health benefited. Friends in Paris soon followed his example and physicians began prescribing the sour-milk diet for their patients.

Bifidobacteria were first isolated from a breast-fed infant by Henry Tissier, who also worked at the Pasteur Institute. The isolated bacterium named Bacillus bifidus communis was later renamed to the genus Bifidobacterium. Tissier found that bifidobacteria are dominant in the gut microbiota of breast-fed babies and he observed clinical benefits from treating diarrhea in infants with bifidobacteria.
During an outbreak of shigellosis in 1917, German professor Alfred Nissle isolated a strain of Escherichia coli from the feces of a soldier who was not affected by the disease. Methods of treating infectious diseases were needed at that time when antibiotics were not yet available, and Nissle used the E. coli Nissle 1917 strain in acute gastrointestinal infectious salmonellosis and shigellosis.

In 1920, Rettger and Cheplin reported that Metchnikoff's "Bulgarian Bacillus", later called Lactobacillus delbrueckii subsp. bulgaricus, could not live in the human intestine. They conducted experiments involving rats and humans volunteers, feeding them with Lactobacillus acidophilus. They observed changes in composition of fecal microbiota, which they described as "transformation of the intestinal flora". Rettger further explored the possibilities of L. acidophilus, and reasoned that bacteria originating from the gut were more likely to produce the desired effect in this environment. In 1935 certain strains of L. acidophilus were found very active when implanted in the human digestive tract. Trials were carried out using this organism, and encouraging results were obtained, especially in the relief of chronic constipation.

Contrasting antibiotics, probiotics were defined as microbially derived factors that stimulate the growth of other microorganisms. In 1989 Roy Fuller suggested a definition of probiotics that has been widely used: "A live microbial feed supplement which beneficially affects the host animal by improving its intestinal microbial balance." Fuller's definition emphasizes the requirement of viability for probiotics and introduces the aspect of a beneficial effect on the host.

The term "probiotic" originally referred to microorganisms that have effects on other microorganisms. The conception of probiotics involved the notion that substances secreted by one microorganism stimulated the growth of another microorganism. The term was used again to describe tissue extracts that stimulated microbial growth. The term probiotics was taken up by Parker, who defined the concept as, "Organisms and substances that have a beneficial effect on the host animal by contributing to its intestinal microbial balance." Later, the definition was greatly improved by Fuller, whose explanation was very close to the definition used today. Fuller described probiotics as a "live microbial feed supplement which beneficially affects the host animal by improving its intestinal microbial balance." He stressed two important claims for probiotics: the viable nature of probiotics and the capacity to help with intestinal balance.

In the following decades, intestinal lactic acid bacterial species with alleged health beneficial properties were introduced as probiotics, including Lactobacillus rhamnosus, Lactobacillus casei, and Lactobacillus johnsonii.

Etymology

Some literature gives the word a full Greek etymology, but it appears to be a composite of the Latin preposition pro, meaning 'for', and the Greek adjective βιωτικός (biōtikos), meaning 'fit for life, lively', the latter deriving from the noun βίος (bios), meaning 'life'. The term contrasts etymologically with the term antibiotic, although it is not a complete antonym. The term probiotic comes from the Latin pro, meaning 'supporting', and refers to a substance that is not digested but "promotes the growth of beneficial intestinal microorganisms".

Research

As food products or dietary supplements, probiotics are under preliminary research to evaluate if they provide any effect on health. In all cases proposed as health claims to the European Food Safety Authority, the scientific evidence remains insufficient to prove a cause-and-effect relationship between consumption of probiotic products and any health benefit. There is no scientific basis for extrapolating an effect from a tested strain to an untested strain. Improved health through gut flora modulation appears to be directly related to long-term dietary changes. According to the National Center for Complementary and Integrative Health: "Although some probiotics have shown promise in research studies, strong scientific evidence to support specific uses of probiotics for most health conditions is lacking."

Claims that some lactobacilli may contribute to weight gain in some humans remain controversial.

Allergies

Probiotics are ineffective in preventing allergies in children, with the possible exception of eczema.

Antibiotic-associated diarrhea

Antibiotics are a common treatment for children, with 11% to 40% of antibiotic-treated children developing diarrhea. Antibiotic-associated diarrhea (AAD) results from an imbalance in the colonic microbiota caused by antibiotic therapy. These microbial community alterations result in changes in carbohydrate metabolism, with decreased short-chain fatty acid absorption and osmotic diarrhea as a result. A 2015 Cochrane review concluded that a protective effect of some probiotics existed for AAD in children. In adults, some probiotics showed a beneficial role in reducing the occurrence of AAD and treating Clostridium difficile disease.

Probiotic treatment might reduce the incidence and severity of AAD as indicated in several meta-analyses. For example, treatment with probiotic formulations including L. rhamnosus may reduce the risk of AAD, improve stool consistency during antibiotic therapy, and enhance the immune response after vaccination.

The potential efficacy of probiotics to treat AAD depends on the probiotic strains and dosage. One review recommended for children L. rhamnosus or Saccharomyces boulardii at 5 to 40 billion colony forming units/day, given the modest number needed to treat and the likelihood that adverse events are very rare. The same review stated that probiotic use should be avoided in pediatric populations at risk for adverse events, such as severely debilitated or immune-compromised children.

Bacterial vaginosis

Probiotic treatment of bacterial vaginosis is the application or ingestion of bacterial species found in the healthy vagina to cure the infection of bacteria causing bacterial vaginosis. This treatment is based on the observation that 70% of healthy females have a group of bacteria in the genus Lactobacillus that dominate the population of organisms in the vagina. Currently, the success of such treatment has been mixed since the use of probiotics to restore healthy populations of Lactobacillus has not been standardized. Often, standard antibiotic treatment is used at the same time that probiotics are being tested. In addition, some groups of women respond to treatment based upon ethnicity, age, number of sexual partners, pregnancy, and the pathogens causing bacterial vaginosis. In 2013 researchers found that administration of hydrogen peroxide producing strains, such as L. acidophilus and L. rhamnosus, were able to normalize vaginal pH and rebalance the vaginal microbiota, preventing and alleviating bacterial vaginosis.

Blood pressure

The consumption of probiotics may modestly help to control high blood pressure.

Cholesterol

Preliminary human and animal studies have demonstrated the efficacy of some strains of lactic acid bacteria (LAB) for reducing serum cholesterol levels, presumably by breaking down bile in the gut, thus inhibiting its reabsorption (where it enters the blood as cholesterol).

A meta-analysis that included five double-blind trials examining the short-term (2–8 weeks) effects of a yogurt with probiotic strains on serum cholesterol levels found a minor change of 8.5 mg/dL (0.22 mmol/L) (4% decrease) in total cholesterol concentration, and a decrease of 7.7 mg/dL (0.2 mmol/L) (5% decrease) in serum LDL concentration.

Diarrhea

Some probiotics are suggested as a possible treatment for various forms of gastroenteritis, and a Cochrane Collaboration meta-analysis on the use of probiotics to treat acute infectious diarrhea based on a comprehensive review of medical literature through 2010 (35 relevant studies, >4500 participants) reported that use of any of the various tested probiotic formulations appeared to reduce the duration of diarrhea by a mean of 25 hours (vs. control groups, 95% confidence interval, 16–34 hours), also noting, however, that "the differences between the studies may be related to other unmeasured and unexplored environmental and host factors" and that further research was needed to confirm reported benefits.

Eczema

Probiotics are commonly given to breast-feeding mothers and their young children to prevent eczema, but some doubt exists over the strength of evidence supporting this effect.

Helicobacter pylori

Some strains of lactic acid bacteria may affect Helicobacter pylori infections (which may cause peptic ulcers) in adults when used in combination with standard medical treatments, but no standard in medical practice or regulatory approval exists for such treatment.

Immune function and infections

Some strains of lactic acid bacteria (LAB) may affect pathogens by means of competitive inhibition (i.e., by competing for growth) and some evidence suggests they may improve immune function by increasing the number of IgA-producing plasma cells and increasing or improving phagocytosis, as well as increasing the proportion of T lymphocytes and natural killer cells. LAB products might aid in the treatment of acute diarrhea, and possibly affect rotavirus infections in children and travelers' diarrhea in adults, but no products are approved for such indications. A large study demonstrated that probiotics may decrease dental caries in children. Two reviews reported reduction of the incidence of respiratory-tract infections in adults.

Probiotics do not appear to change the risk of infection in older people.

Inflammatory bowel disease

Probiotics are being studied for their potential to influence inflammatory bowel disease. There is some evidence to support their use in conjunction with standard medications in treating ulcerative colitis and no evidence of their efficacy in treating Crohn's disease.

A live formulation of lyophilized Bifidobacterium breve, Bifidobacterium longum, Bifidobacterium infantis, Lactobacillus acidophilus, Lactobacillus plantarum, Lactobacillus paracasei, Lactobacillus bulgaricus, and Streptococcus thermophilus (VSL#3) has shown effectiveness in the small clinical trials, some of which were not randomized nor double-blinded, that had been done as of 2015; more high-quality clinical trials are needed to determine safety and effectiveness.

Irritable bowel syndrome

Probiotics are under study for their potential to affect irritable bowel syndrome, although uncertainty remains around which type of probiotic works best, and around the size of possible effect.

Lactose intolerance

Ingestion of certain active strains may help lactose-intolerant individuals tolerate more lactose than they would otherwise have tolerated.

Necrotizing enterocolitis

Several clinical studies provide evidence for the potential of probiotics to lower the risk of necrotizing enterocolitis and mortality in premature infants. One meta-analysis indicated that probiotics reduce these risks by more than 50% compared with controls.

Recurrent abdominal pain

A 2017 review based on moderate to low-quality evidences suggests that probiotics may be helpful in relieving pain in the short term in children with recurrent abdominal pain, but the proper strain and dosage are not known.

Urinary tract

There is no good evidence that probiotics are of benefit in the management of infection or inflammation of the urinary tract.

General research

Formulations

Supplements such as tablets, capsules, powders, and sachets containing the bacteria have been studied. However, probiotics taken orally can be destroyed by the acidic conditions of the stomach. As of 2010, a number of microencapsulation techniques were being developed to address this problem.

Multiple probiotics

Preliminary research is evaluating the potential physiological effects of multiple probiotic strains, as opposed to a single strain. As the human gut may contain several hundred microbial species, one theory indicates that this diverse environment may benefit from consuming multiple probiotic strains, an effect that remains scientifically unconfirmed.

Strains

There is only preliminary evidence for most probiotic health claims. Even for the most studied probiotic strains, few have been sufficiently developed in basic and clinical research to warrant approval for health claim status by a regulatory agency such as the Food and Drug Administration or European Food Safety Authority, and, as of 2010, no claims had been approved by those two agencies. Some experts are skeptical about the efficacy of different probiotic strains and believe that not all subjects benefit from probiotics.

Scientific guidelines for testing

First, probiotics must be alive when administered. One of the concerns throughout the scientific literature resides in the viability and reproducibility on a large scale of observed results for specific studies, as well as the viability and stability during use and storage, and finally the ability to survive in stomach acids and then in the intestinal ecosystem.

Secondly, probiotics must have undergone controlled evaluation to document health benefits in the target host. Only products that contain live organisms shown in reproducible human studies to confer a health benefit can actually claim to be probiotic. The correct definition of health benefit, backed with solid scientific evidence, is a strong element for the proper identification and assessment of the effect of a probiotic. This aspect represents a major challenge for scientific and industrial investigations because several difficulties arise, such as variability in the site for probiotic use (oral, vaginal, intestinal) and mode of application.

Thirdly, the probiotic candidate must be a taxonomically defined microbe or combination of microbes (genus, species, and strain level). It is commonly admitted that most effects of probiotics are strain-specific and cannot be extended to other probiotics of the same genus or species. This calls for a precise identification of the strain, i.e. genotypic and phenotypic characterization of the tested microorganism.

Fourthly, probiotics must be safe for their intended use. The 2002 FAO/WHO guidelines recommend that, though bacteria may be generally recognized as safe (GRAS), the safety of the potential probiotic should be assessed by the minimum required tests:

• Determination of antibiotic resistance patterns
• Assessment of certain metabolic activities (e.g. D-lactate production, bile salt deconjugation)
• Assessment of side effects during human studies
• Epidemiological surveillance of adverse incidents in consumers (after market)
• If the strain under evaluation belongs to a species that is a known mammalian toxin producer, it must be tested for toxin production. One possible scheme for testing toxin production has been recommended by the EU Scientific Committee on Animal Nutrition.
• If the strain under evaluation belongs to a species with known hemolytic potential, determination of hemolytic activity is required.

In Europe, the EFSA has adopted a premarket system for safety assessment of microbial species used in food and feed productions to set priorities for the need of risk assessment. The assessment is made for a selected group of microorganisms, which if favorable, leads to a “Qualified Presumption of Safety” status.

Fifthly and finally, probiotics must be supplied in adequate numbers, which may be defined as the number able to trigger the targeted effect on the host. It depends on strain specificity, process, and matrix, as well as the targeted effect. Most of the reported benefits demonstrated with the traditional probiotics have been observed after ingestion of a concentration around 10⁷ to 10⁸ probiotic cells per gram, with a serving size around 100 to 200 mg per day.

Microbiota

From Wikipedia, the free encyclopedia

The predominant bacteria on human skin

A microbiota is an "ecological community of commensal, symbiotic and pathogenic microorganisms" found in and on all multicellular organisms studied to date from plants to animals. A microbiota includes bacteria, archaea, protists, fungi and viruses. Microbiota have been found to be crucial for immunologic, hormonal and metabolic homeostasis of their host. The synonymous term microbiome describes either the collective genomes of the microorganisms that reside in an environmental niche or the microorganisms themselves.

The microbiome and host emerged during evolution as a synergistic unit from epigenetics and genetic characteristics, sometimes collectively referred to as a holobiont.

Introduction

All plants and animals, from simple life forms to humans, live in close association with microbial organisms. Several advances have driven the perception of microbiomes, including:

• the ability to perform genomic and gene expression analyses of single cells and of entire microbial communities in the disciplines of metagenomics and metatranscriptomics
• databases accessible to researchers across multiple disciplines
• methods of mathematical analysis suitable for complex data sets

Biologists have come to appreciate that microbes make up an important part of an organism's phenotype, far beyond the occasional symbiotic case study.

Types of host relationships

Commensalism, a concept developed by Pierre-Joseph van Beneden (1809-1894), a Belgian professor at the University of Louvain during the nineteenth century is central to the microbiome, where microbiota colonize a host in a non-harmful coexistence. The relationship with their host is called mutualistic when organisms perform tasks that are known to be useful for the host, parasitic, when disadvantageous to the host. Other authors define a situation as mutualistic where both benefit, and commensal, where the unaffected host benefits the symbiont. A nutrient exchange may be bidirectional or unidirectional, may be context dependent and may occur in diverse ways. Microbiota that are expected to be present, and that under normal circumstances do not cause disease, are deemed normal flora or normal microbiota.

Acquisition and change

The initial acquisition of microbiota in animals from mammalians to marine sponges is at birth, and may even occur through the germ cell line. In plants, the colonizing process can be initiated below ground in the root zone, around the germinating seed, the spermosphere, or originate from the above ground parts, the phyllosphere and the flower zone or anthosphere. The stability of the rhizosphere microbiota over generations depends upon the plant type but even more on the soil composition, i.e. living and non living environment.

Microbiota by host

Consensus exists among evolutionary biologists that one should not separate an organism's genes from the context of its resident microbes.

Humans

The human microbiota includes bacteria, fungi, archaea and viruses. Micro-animals which live on the human body are excluded. The human microbiome refers to their genomes.

Humans are colonized by many microorganisms; the traditional estimate was that humans live with ten times more non-human cells than human cells; more recent estimates have lowered this to 3:1 and even to about 1:1.

The Human Microbiome Project sequenced the genome of the human microbiota, focusing particularly on the microbiota that normally inhabit the skin, mouth, nose, digestive tract, and vagina. It reached a milestone in 2012 when it published initial results.

Non-human animals

• Amphibians have microbiota on their skin. Some species are able to carry a fungus named Batrachochytrium dendrobatidis, which in others can cause a deadly infection Chytridiomycosis depending on their microbiome, resisting pathogen colonization or inhibiting their growth with antimicrobial skin peptides.
• In mammals, herbivores such as cattle depend on their rumen microbiome to convert cellulose into proteins, short chain fatty acids, and gases. Culture methods cannot provide information on all microorganisms present. Comparative metagenomic studies yielded the surprising result that individual cattle possess markedly different community structures, predicted phenotype, and metabolic potentials, even though they were fed identical diets, were housed together, and were apparently functionally identical in their utilization of plant cell wall resources.
• Mice have become the most studied mammalian regarding their microbiomes. The gut microbiota have been studied in relation to allergic airway disease, obesity, gastrointestinal diseases and diabetes. Perinatal shifting of microbiota through low dose antibiotics can have long-lasting effects on future susceptibility to allergic airway disease. The frequency of certain subsets of microbes has been linked to disease severity. The presence of specific microbes early in postnatal life, instruct future immune responses. In gnotobiotic mice certain gut bacteria were found to transmit a particular phenotype to recipient germ-free mice, that promoted accumulation of colonic regulatory T cells, and strains that modulated mouse adiposity and cecal metabolite concentrations. This combinatorial approach enables a systems-level understanding of microbial contributions to human biology. But also other mucoide tissues as lung and vagina have been studied in relation to diseases such as asthma, allergy and vaginosis.
• Insects have their own microbiomes. For example, leaf-cutter ants form huge underground colonies harvesting hundreds of kilograms of leaves each year and are unable to digest the cellulose in the leaves directly. They maintain fungus gardens as the colony's primary food source. While the fungus itself does not digest cellulose, a microbial community containing a diversity of bacteria is doing so. Analysis of the microbial population's genome revealed many genes with a role in cellulose digestion. This microbiome's predicted carbohydrate-degrading enzyme profile is similar to that of the bovine rumen, but the species composition is almost entirely different. Gut microbiota of the fruit fly can affect the way its gut looks, by impacting epithelial renewal rate, cellular spacing, and the composition of different cell types in the epithelium. When the moth Spodoptera exigua is infected with baculovirus immune-related genes are downregulated and the amount of its gut microbiota increases.

Plants

Light micrograph of a cross section of a coralloid root of a cycad, showing the layer that hosts symbiotic cyanobacteria

• Plants are attractive hosts for microorganisms since they provide a variety of nutrients. Microorganisms on plants can be epiphytes (found on the plants) or endophytes (found inside plant tissue). Oomycetes and fungi have, through convergent evolution, developed similar morphology and occupy similar ecological niches. They develop hyphae, threadlike structures that penetrate the host cell. In mutualistic situations the plant often exchanges hexose sugars for inorganic phosphate from the fungal symbiont. It is speculated that such very ancient associations have aided plants when they first colonized land. Plant-growth promoting bacteria (PGPB) provide the plant with essential services such as nitrogen fixation, solubilization of minerals such as phosphorus, synthesis of plant hormones, direct enhancement of mineral uptake, and protection from pathogens. PGPBs may protect plants from pathogens by competing with the pathogen for an ecological niche or a substrate, producing inhibitory allelochemicals, or inducing systemic resistance in host plants to the pathogen

Functions

Microbiota have been found to be crucial for immunologic, hormonal and metabolic homeostasis of their host.

Immune system

The symbiotic relationship between a host and its microbiota shapes the immune system of mammalians, insects, plants and aquatic organisms. In many animals, the immune system and microbiota engage in "cross-talk", exchanging chemical signals. This allows the immune system to recognize the types of bacteria that are harmful to the host and combat them, while allowing the helpful bacteria to carry out their functions; in turn, the microbiota influence immune reactivity and targeting. Bacteria can be transferred from mother to child through direct contact and after birth, or through indirect contact through eggs, coprophagy, and several other pathways. As the infant microbiome is established, commensal bacteria quickly populate the gut, prompting a range of immune responses and "programming" the immune system with long-lasting effects. The bacteria are able to stimulate lymphoid tissue associated with the gut mucosa, which enables the tissue to produce antibodies for pathogens that may enter the gut.

The human microbiome plays a role in the activation of toll-like receptors (TLRs) in the intestines, a type of pattern recognition receptor host cells use to recognize dangers and repair damage. Pathogens can influence this coexistence leading to immune dysregulation including and susceptibility to diseases, immune tolerance and autoimmune diseases.

Endocrine system

Intestinal microbiota can interact with thyroid-related micronutrients and the metabolism of endogenous iodothyronines such as Triiodothyronine and exogenous iodothyronines, which affects even phyla without thyroid follicles, such as jellyfish, insects, and sea urchins, as iodothyronine-induced metamorphosis is an ancestral feature of all chordates.

Microbiota can affect a fruit fly´s sex preference in mating. Infecting flies with pure cultures of Lactobacillus plantarum established a certain mating preference as L. plantarum can change the fly´s levels of cuticular hydrocarbon sex pheromones.

Metabolism

When adult germ-free mice are colonized with the gut flora of obese mice, they can gain weight dramatically with an increased metabolism of monosaccharides and short-chain fatty acids. The gut flora of obese mice has less Bacteroidetes than Firmicutes and is thought to be more efficient at extracting energy from food.

Co-evolution of microbiota

Bleached branching coral (foreground) and normal branching coral (background). Keppel Islands, Great Barrier Reef

Organisms evolve within eco-systems so that the change of one organism affects the change of others. Co-evolution (also called "hologenome theory") proposes that an object of natural selection is not the individual organism, but the organism together with its associated organisms, including its microbial communities.

Coral reefs. The hologenome theory originated in studies on coral reefs. Coral reefs are the largest structures created by living organisms, and contain abundant and highly complex microbial communities. Over the past several decades, major declines in coral populations have occurred. Climate change, water pollution and over-fishing are three stress factors that have been described as leading to disease susceptibility. Over twenty different coral diseases have been described, but of these, only a handful have had their causative agents isolated and characterized. Coral bleaching is the most serious of these diseases. In the Mediterranean Sea, the bleaching of Oculina patagonica was first described in 1994 and shortly determined to be due to infection by Vibrio shiloi. From 1994 to 2002, bacterial bleaching of O. patagonica occurred every summer in the eastern Mediterranean. Surprisingly, however, after 2003, O. patagonica in the eastern Mediterranean has been resistant to V. shiloi infection, although other diseases still cause bleaching. The surprise stems from the knowledge that corals are long lived, with lifespans on the order of decades, and do not have adaptive immune systems. Their innate immune systems do not produce antibodies, and they should seemingly not be able to respond to new challenges except over evolutionary time scales.

The puzzle of how corals managed to acquire resistance to a specific pathogen led to a 2007 proposal, that a dynamic relationship exists between corals and their symbiotic microbial communities. It is thought that by altering its composition, the holobiont can adapt to changing environmental conditions far more rapidly than by genetic mutation and selection alone. Extrapolating this hypothesis to other organisms, including higher plants and animals, led to the proposal of the "hologenome theory of evolution".

As of 2007 the hologenome theory was still being debated. A major criticism has been the claim that V. shiloi was misidentified as the causative agent of coral bleaching, and that its presence in bleached O. patagonica was simply that of opportunistic colonization. If this is true, the basic observation leading to the theory would be invalid. The theory has gained significant popularity as a way of explaining rapid changes in adaptation that cannot otherwise be explained by traditional mechanisms of natural selection. Within the hologenome theory, the holobiont has not only become the principal unit of natural selection but also the result of other step of integration that it is also observed at the cell (symbiogenesis, endosymbiosis) and genomic levels.

Research methods

Targeted amplicon sequencing

Targeted amplicon sequencing relies on having some expectations about the composition of the community that is being studied. In target amplicon sequencing a phylogenetically informative marker is targeted for sequencing. Such a marker should be present in ideally all the expected organisms. It should also evolve in such a way that it is conserved enough that primers can target genes from a wide range of organisms while evolving quickly enough to allow for finer resolution at the taxonomic level. A common marker for human microbiome studies is the gene for bacterial 16S rRNA (i.e. "16S rDNA", the sequence of DNA which encodes the ribosomal RNA molecule). Since ribosomes are present in all living organisms, using 16S rDNA allows for DNA to be amplified from many more organisms than if another marker were used. The 16S rDNA gene contains both slowly evolving regions and fast evolving regions; the former can be used to design broad primers while the latter allow for finer taxonomic distinction. However, species-level resolution is not typically possible using the 16S rDNA. Primer selection is an important step, as anything that cannot be targeted by the primer will not be amplified and thus will not be detected. Different sets of primers have been shown to amplify different taxonomic groups due to sequence variation.

Targeted studies of eukaryotic and viral communities are limited and subject to the challenge of excluding host DNA from amplification and the reduced eukaryotic and viral biomass in the human microbiome.

After the amplicons are sequenced, molecular phylogenetic methods are used to infer the composition of the microbial community. This is done by clustering the amplicons into operational taxonomic units (OTUs) and inferring phylogenetic relationships between the sequences. Due to the complexity of the data, distance measures such as UniFrac distances are usually defined between microbiome samples, and downstream multivariate methods are carried out on the distance matrices. An important point is that the scale of data is extensive, and further approaches must be taken to identify patterns from the available information. Tools used to analyze the data include VAMPS, QIIME and mothur.

Metagenomic sequencing

Metagenomics is also used extensively for studying microbial communities. In metagenomic sequencing, DNA is recovered directly from environmental samples in an untargeted manner with the goal of obtaining an unbiased sample from all genes of all members of the community. Recent studies use shotgun Sanger sequencing or pyrosequencing to recover the sequences of the reads. The reads can then be assembled into contigs. To determine the phylogenetic identity of a sequence, it is compared to available full genome sequences using methods such as BLAST. One drawback of this approach is that many members of microbial communities do not have a representative sequenced genome.

Despite the fact that metagenomics is limited by the availability of reference sequences, one significant advantage of metagenomics over targeted amplicon sequencing is that metagenomics data can elucidate the functional potential of the community DNA. Targeted gene surveys cannot do this as they only reveal the phylogenetic relationship between the same gene from different organisms. Functional analysis is done by comparing the recovered sequences to databases of metagenomic annotations such as KEGG. The metabolic pathways that these genes are involved in can then be predicted with tools such as MG-RAST, CAMERA, and IMG/M.

RNA and protein-based approaches

Metatranscriptomics studies have been performed to study the gene expression of microbial communities through methods such as the pyrosequencing of extracted RNA. Structure based studies have also identified non-coding RNAs (ncRNAs) such as ribozymes from microbiota.  Metaproteomics is an approach that studies the proteins expressed by microbiota, giving insight into its functional potential.

Projects

The Human Microbiome Project launched in 2008 was a United States National Institutes of Health initiative to identify and characterize microorganisms found in both healthy and diseased humans. The five-year project, best characterized as a feasibility study with a budget of $115 million tested how changes in the human microbiome are associated with human health or disease.

The Earth Microbiome Project (EMP) is an initiative to collect natural samples and analyze the microbial community around the globe. Microbes are highly abundant, diverse and have an important role in the ecological system. Yet as of 2010, it was estimated that the total global environmental DNA sequencing effort had produced less than 1 percent of the total DNA found in a liter of seawater or a gram of soil, and the specific interactions between microbes are largely unknown. The EMP aims to process as many as 200,000 samples in different biomes, generating a complete database of microbes on earth to characterize environments and ecosystems by microbial composition and interaction. Using these data, new ecological and evolutionary theories can be proposed and tested.

The Brazilian Microbiome Project aims to assemble a Brazilian Microbiome Consortium/Database. This is the first attempt to collect and collate information about Brazilian microbial genetic and functional diversity in a systematic and holistic manner. New sequence data have been generated from samples collected in all Brazilian regions.

Privacy issues

Microbial DNA inhabiting a person's human body can uniquely identify the person. A person's privacy may be compromised if the person anonymously donated microbe DNA data. Their medical condition and identity could be revealed.

Starch

From Wikipedia, the free encyclopedia

Starch

Identifiers
CAS Number	9005-25-8
ChemSpider	none
ECHA InfoCard	100.029.696
EC Number	232-679-6
RTECS number	GM5090000
Properties
Chemical formula	(C 6H 10O 5) n - (H 2O)
Molar mass	Variable
Appearance	White powder
Density	Variable
Melting point	decomposes
Solubility in water	insoluble
Thermochemistry
Std enthalpy of combustion (ΔcHo298)	4.1788 kilocalories per gram (17.484 kJ/g)
Hazards
Safety data sheet	ICSC 1553
Autoignition temperature	410 °C (770 °F; 683 K)
US health exposure limits (NIOSH):
PEL (Permissible)	TWA 15 mg/m3 (total) TWA 5 mg/m3 (resp)
Except where otherwise noted, data are given for materials in their standard state (at 25 °C [77 °F], 100 kPa).

Structure of the amylose molecule

 

Structure of the amylopectin molecule

Starch or amylum is a polymeric carbohydrate consisting of a large number of glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants as energy storage. It is the most common carbohydrate in human diets and is contained in large amounts in staple foods like potatoes, wheat, maize (corn), rice, and cassava.

Pure starch is a white, tasteless and odorless powder that is insoluble in cold water or alcohol. It consists of two types of molecules: the linear and helical amylose and the branched amylopectin. Depending on the plant, starch generally contains 20 to 25% amylose and 75 to 80% amylopectin by weight. Glycogen, the glucose store of animals, is a more highly branched version of amylopectin.

In industry, starch is converted into sugars, for example by malting, and fermented to produce ethanol in the manufacture of beer, whisky and biofuel. It is processed to produce many of the sugars used in processed foods. Mixing most starches in warm water produces a paste, such as wheatpaste, which can be used as a thickening, stiffening or gluing agent. The biggest industrial non-food use of starch is as an adhesive in the papermaking process. Starch can be applied to parts of some garments before ironing, to stiffen them.

Etymology

The word "starch" is from a Germanic root with the meanings "strong, stiff, strengthen, stiffen". Modern German Stärke (strength) is related. The Greek term for starch, "amylon" (ἄμυλον), is also related. It provides the root amyl, which is used as a prefix in biochemistry for several 5-carbon compounds related to or derived from starch (e.g. amyl alcohol).

History

Starch grains from the rhizomes of Typha (cattails, bullrushes) as flour have been identified from grinding stones in Europe dating back to 30,000 years ago. Starch grains from sorghum were found on grind stones in caves in Ngalue, Mozambique dating up to 100,000 years ago.

Pure extracted wheat starch paste was used in Ancient Egypt possibly to glue papyrus. The extraction of starch is first described in the Natural History of Pliny the Elder around AD 77–79. Romans used it also in cosmetic creams, to powder the hair and to thicken sauces. Persians and Indians used it to make dishes similar to gothumai wheat halva. Rice starch as surface treatment of paper has been used in paper production in China since 700 CE.

In addition to starchy plants consumed directly, 66 million tonnes of starch were being produced per year worldwide by 2008. In the EU this was around 8.5 million tonnes, with around 40% being used for industrial applications and 60% for food uses, most of the latter as glucose syrups.

Energy store of plants

Most green plants use starch as their energy store.The extra glucose is changed into starch which is more complex than glucose(by plants). An exception is the family Asteraceae (asters, daisies and sunflowers), where starch is replaced by the fructan inulin. Inulin-like fructans are also present in grasses such as wheat, in onions and garlic, bananas, and asparagus.

In photosynthesis, plants use light energy to produce glucose from carbon dioxide. The glucose is used to generate the chemical energy required for general metabolism, to make organic compounds such as nucleic acids, lipids, proteins and structural polysaccharides such as cellulose, or is stored in the form of starch granules, in amyloplasts. Toward the end of the growing season, starch accumulates in twigs of trees near the buds. Fruit, seeds, rhizomes, and tubers store starch to prepare for the next growing season.

Glucose is soluble in water, hydrophilic, binds with water and then takes up much space and is osmotically active; glucose in the form of starch, on the other hand, is not soluble, therefore osmotically inactive and can be stored much more compactly.

Glucose molecules are bound in starch by the easily hydrolyzed alpha bonds. The same type of bond is found in the animal reserve polysaccharide glycogen. This is in contrast to many structural polysaccharides such as chitin, cellulose and peptidoglycan, which are bound by beta bonds and are much more resistant to hydrolysis.

Biosynthesis

Plants produce starch by first converting glucose 1-phosphate to ADP-glucose using the enzyme glucose-1-phosphate adenylyltransferase. This step requires energy in the form of ATP. The enzyme starch synthase then adds the ADP-glucose via a 1,4-alpha glycosidic bond to a growing chain of glucose residues, liberating ADP and creating amylose. The ADP-glucose is almost certainly added to the non-reducing end of the amylose polymer, as the UDP-glucose is added to the non-reducing end of glycogen during glycogen synthesis.

Starch branching enzyme introduces 1,6-alpha glycosidic bonds between the amylose chains, creating the branched amylopectin. The starch debranching enzyme isoamylase removes some of these branches. Several isoforms of these enzymes exist, leading to a highly complex synthesis process.

Glycogen and amylopectin have similar structure, but the former has about one branch point per ten 1,4-alpha bonds, compared to about one branch point per thirty 1,4-alpha bonds in amylopectin. Amylopectin is synthesized from ADP-glucose while mammals and fungi synthesize glycogen from UDP-glucose; for most cases, bacteria synthesize glycogen from ADP-glucose (analogous to starch).

In addition to starch synthesis in plants, starch can be synthesized from non-food starch mediated by an enzyme cocktail. In this cell-free biosystem, beta-1,4-glycosidic bond-linked cellulose is partially hydrolyzed to cellobiose. Cellobiose phosphorylase cleaves to glucose 1-phosphate and glucose; the other enzyme—potato alpha-glucan phosphorylase can add a glucose unit from glucose 1-phosphorylase to the non-reducing ends of starch. In it, phosphate is internally recycled. The other product, glucose, can be assimilated by a yeast. This cell-free bioprocessing does not need any costly chemical and energy input, can be conducted in aqueous solution, and does not have sugar losses.

Degradation

Starch is synthesized in plant leaves during the day and stored as granules; it serves as an energy source at night. The insoluble, highly branched starch chains have to be phosphorylated in order to be accessible for degrading enzymes. The enzyme glucan, water dikinase (GWD) phosphorylates at the C-6 position of a glucose molecule, close to the chains 1,6-alpha branching bonds. A second enzyme, phosphoglucan, water dikinase (PWD) phosphorylates the glucose molecule at the C-3 position. A loss of these enzymes, for example a loss of the GWD, leads to a starch excess (sex) phenotype, and because starch cannot be phosphorylated, it accumulates in the plastids.

After the phosphorylation, the first degrading enzyme, beta-amylase (BAM) can attack the glucose chain at its non-reducing end. Maltose is released as the main product of starch degradation. If the glucose chain consists of three or fewer molecules, BAM cannot release maltose. A second enzyme, disproportionating enzyme-1 (DPE1), combines two maltotriose molecules. From this chain, a glucose molecule is released. Now, BAM can release another maltose molecule from the remaining chain. This cycle repeats until starch is degraded completely. If BAM comes close to the phosphorylated branching point of the glucose chain, it can no longer release maltose. In order for the phosphorylated chain to be degraded, the enzyme isoamylase (ISA) is required.

The products of starch degradation are predominantly maltose and smaller amounts of glucose. These molecules are exported from the plastid to the cytosol, maltose via the maltose transporter, which if mutated (MEX1-mutant) results in maltose accumulation in the plastid. Glucose is exported via the plastidic glucose translocator (pGlcT). These two sugars act as a precursor for sucrose synthesis.  Sucrose can then be used in the oxidative pentose phosphate pathway in the mitochondria, to generate ATP at night.

Properties

Structure

Starch, 800x magnified, under polarized light, showing characteristic extinction cross

 

Rice starch seen on light microscope. Characteristic for the rice starch is that starch granules have an angular outline and some of them are attached to each other and form larger granules

While amylose was thought to be completely unbranched, it is now known that some of its molecules contain a few branch points. Amylose is a much smaller molecule than amylopectin. About one quarter of the mass of starch granules in plants consist of amylose, although there are about 150 times more amylose than amylopectin molecules.

Starch molecules arrange themselves in the plant in semi-crystalline granules. Each plant species has a unique starch granular size: rice starch is relatively small (about 2 μm) while potato starches have larger granules (up to 100 μm).

Starch becomes soluble in water when heated. The granules swell and burst, the semi-crystalline structure is lost and the smaller amylose molecules start leaching out of the granule, forming a network that holds water and increasing the mixture's viscosity. This process is called starch gelatinization. During cooking, the starch becomes a paste and increases further in viscosity. During cooling or prolonged storage of the paste, the semi-crystalline structure partially recovers and the starch paste thickens, expelling water. This is mainly caused by retrogradation of the amylose. This process is responsible for the hardening of bread or staling, and for the water layer on top of a starch gel (syneresis).

Some cultivated plant varieties have pure amylopectin starch without amylose, known as waxy starches. The most used is waxy maize, others are glutinous rice and waxy potato starch. Waxy starches have less retrogradation, resulting in a more stable paste. High amylose starch, amylomaize, is cultivated for the use of its gel strength and for use as a resistant starch (a starch that resists digestion) in food products.

Synthetic amylose made from cellulose has a well-controlled degree of polymerization. Therefore, it can be used as a potential drug deliver carrier.

Certain starches, when mixed with water, will produce a non-newtonian fluid sometimes nicknamed "oobleck".

Hydrolysis

The enzymes that break down or hydrolyze starch into the constituent sugars are known as amylases.
Alpha-amylases are found in plants and in animals. Human saliva is rich in amylase, and the pancreas also secretes the enzyme. Individuals from populations with a high-starch diet tend to have more amylase genes than those with low-starch diets.

Beta-amylase cuts starch into maltose units. This process is important in the digestion of starch and is also used in brewing, where amylase from the skin of seed grains is responsible for converting starch to maltose (Malting, Mashing).

Given a heat of combustion of glucose of 2,805 kilojoules per mole (670 kcal/mol) whereas that of starch is 2,835 kJ (678 kcal) per mole of glucose monomer, hydrolysis releases about 30 kJ (7.2 kcal) per mole, or 166 J (40 cal) per gram of glucose product.

Dextrinization

If starch is subjected to dry heat, it breaks down to form dextrins, also called "pyrodextrins" in this context. This break down process is known as dextrinization. (Pyro)dextrins are mainly yellow to brown in color and dextrinization is partially responsible for the browning of toasted bread.

Chemical tests

Granules of wheat starch, stained with iodine, photographed through a light microscope

A triiodide (I₃⁻) solution formed by mixing iodine and iodide (usually from potassium iodide) is used to test for starch; a dark blue color indicates the presence of starch. The details of this reaction are not fully known, but recent scientific work using single crystal x-ray crystallography and comparative Raman spectroscopy suggests that the final starch-iodine structure is similar to an infinite polyiodide chain like one found in a pyrroloperylene-iodine complex. The strength of the resulting blue color depends on the amount of amylose present. Waxy starches with little or no amylose present will color red. Benedict's test and Fehling's test is also done to indicate the presence of starch.

Starch indicator solution consisting of water, starch and iodide is often used in redox titrations: in the presence of an oxidizing agent the solution turns blue, in the presence of reducing agent the blue color disappears because triiodide (I₃⁻) ions break up into three iodide ions, disassembling the starch-iodine complex. Starch solution was used as indicator for visualizing the periodic formation and consumption of triiodide intermediate in the Briggs-Rauscher oscillating reaction. The starch, however, changes the kinetics of the reaction steps involving triiodide ion. A 0.3% w/w solution is the standard concentration for a starch indicator. It is made by adding 3 grams of soluble starch to 1 liter of heated water; the solution is cooled before use (starch-iodine complex becomes unstable at temperatures above 35 °C).

Each species of plant has a unique type of starch granules in granular size, shape and crystallization pattern. Under the microscope, starch grains stained with iodine illuminated from behind with polarized light show a distinctive Maltese cross effect (also known as extinction cross and birefringence).

Food

Starch is the most common carbohydrate in the human diet and is contained in many staple foods. The major sources of starch intake worldwide are the cereals (rice, wheat, and maize) and the root vegetables (potatoes and cassava). Many other starchy foods are grown, some only in specific climates, including acorns, arrowroot, arracacha, bananas, barley, breadfruit, buckwheat, canna, colacasia, katakuri, kudzu, malanga, millet, oats, oca, polynesian arrowroot, sago, sorghum, sweet potatoes, rye, taro, chestnuts, water chestnuts and yams, and many kinds of beans, such as favas, lentils, mung beans, peas, and chickpeas.

Widely used prepared foods containing starch are bread, pancakes, cereals, noodles, pasta, porridge and tortilla.

Digestive enzymes have problems digesting crystalline structures. Raw starch is digested poorly in the duodenum and small intestine, while bacterial degradation takes place mainly in the colon. When starch is cooked, the digestibility is increased.

Starch gelatinization during cake baking can be impaired by sugar competing for water, preventing gelatinization and improving texture.

Before the advent of processed foods, people consumed large amounts of uncooked and unprocessed starch-containing plants, which contained high amounts of resistant starch. Microbes within the large intestine fermented the starch, produced short-chain fatty acids, which are used as energy, and support the maintenance and growth of the microbes. More highly processed foods are more easily digested and release more glucose in the small intestine—less starch reaches the large intestine and more energy is absorbed by the body. It is thought that this shift in energy delivery (as a result of eating more processed foods) may be one of the contributing factors to the development of metabolic disorders of modern life, including obesity and diabetes.

Starch industry

The starch industry extracts and refines starches from seeds, roots and tubers, by wet grinding, washing, sieving and drying. Today, the main commercial refined starches are cornstarch, tapioca, arrowroot, and wheat, rice, and potato starches. To a lesser extent, sources of refined starch are sweet potato, sago and mung bean. To this day, starch is extracted from more than 50 types of plants.

Untreated starch requires heat to thicken or gelatinize. When a starch is pre-cooked, it can then be used to thicken instantly in cold water. This is referred to as a pregelatinized starch.

Starch sugars

Starch can be hydrolyzed into simpler carbohydrates by acids, various enzymes, or a combination of the two. The resulting fragments are known as dextrins. The extent of conversion is typically quantified by dextrose equivalent (DE), which is roughly the fraction of the glycosidic bonds in starch that have been broken.

These starch sugars are by far the most common starch based food ingredient and are used as sweeteners in many drinks and foods. They include:

• Maltodextrin, a lightly hydrolyzed (DE 10–20) starch product used as a bland-tasting filler and thickener.
• Various glucose syrups (DE 30–70), also called corn syrups in the US, viscous solutions used as sweeteners and thickeners in many kinds of processed foods.
• Dextrose (DE 100), commercial glucose, prepared by the complete hydrolysis of starch.
• High fructose syrup, made by treating dextrose solutions with the enzyme glucose isomerase, until a substantial fraction of the glucose has been converted to fructose. In the United States sugar prices are two to three times higher than in the rest of the world; high-fructose corn syrup is significantly cheaper, and is the principal sweetener used in processed foods and beverages. Fructose also has better microbiological stability. One kind of high fructose corn syrup, HFCS-55, is sweeter than sucrose because it is made with more fructose, while the sweetness of HFCS-42 is on par with sucrose.
• Sugar alcohols, such as maltitol, erythritol, sorbitol, mannitol and hydrogenated starch hydrolysate, are sweeteners made by reducing sugars.

Modified starches

A modified starch is a starch that has been chemically modified to allow the starch to function properly under conditions frequently encountered during processing or storage, such as high heat, high shear, low pH, freeze/thaw and cooling.

The modified food starches are E coded according to the International Numbering System for Food Additives (INS):

• 1400 Dextrin
• 1401 Acid-treated starch
• 1402 Alkaline-treated starch
• 1403 Bleached starch
• 1404 Oxidized starch
• 1405 Starches, enzyme-treated
• 1410 Monostarch phosphate
• 1412 Distarch phosphate
• 1413 Phosphated distarch phosphate
• 1414 Acetylated distarch phosphate
• 1420 Starch acetate
• 1422 Acetylated distarch adipate
• 1440 Hydroxypropyl starch
• 1442 Hydroxypropyl distarch phosphate
• 1443 Hydroxypropyl distarch glycerol
• 1450 Starch sodium octenyl succinate
• 1451 Acetylated oxidized starch

INS 1400, 1401, 1402, 1403 and 1405 are in the EU food ingredients without an E-number. Typical modified starches for technical applications are cationic starches, hydroxyethyl starch and carboxymethylated starches.

Use as food additive

As an additive for food processing, food starches are typically used as thickeners and stabilizers in foods such as puddings, custards, soups, sauces, gravies, pie fillings, and salad dressings, and to make noodles and pastas. Function as thickeners, extenders, emulsion stabilizers and are exceptional binders in processed meats.

Gummed sweets such as jelly beans and wine gums are not manufactured using a mold in the conventional sense. A tray is filled with native starch and leveled. A positive mold is then pressed into the starch leaving an impression of 1,000 or so jelly beans. The jelly mix is then poured into the impressions and put onto a stove to set. This method greatly reduces the number of molds that must be manufactured.

Use in pharmaceutical industry

In the pharmaceutical industry, starch is also used as an excipient, as tablet disintegrant, and as binder.

Resistant starch

Resistant starch is starch that escapes digestion in the small intestine of healthy individuals. High amylose starch from corn has a higher gelatinization temperature than other types of starch and retains its resistant starch content through baking, mild extrusion and other food processing techniques. It is used as an insoluble dietary fiber in processed foods such as bread, pasta, cookies, crackers, pretzels and other low moisture foods. It is also utilized as a dietary supplement for its health benefits. Published studies have shown that resistant starch helps to improve insulin sensitivity, increases satiety and improves markers of colonic function. It has been suggested that resistant starch contributes to the health benefits of intact whole grains.

Industrial applications

Starch adhesive

Papermaking

Papermaking is the largest non-food application for starches globally, consuming millions of metric tons annually. In a typical sheet of copy paper for instance, the starch content may be as high as 8%. Both chemically modified and unmodified starches are used in papermaking. In the wet part of the papermaking process, generally called the "wet-end", the starches used are cationic and have a positive charge bound to the starch polymer. These starch derivatives associate with the anionic or negatively charged paper fibers / cellulose and inorganic fillers. Cationic starches together with other retention and internal sizing agents help to give the necessary strength properties to the paper web formed in the papermaking process (wet strength), and to provide strength to the final paper sheet (dry strength).

In the dry end of the papermaking process, the paper web is rewetted with a starch based solution. The process is called surface sizing. Starches used have been chemically, or enzymatically depolymerized at the paper mill or by the starch industry (oxidized starch). The size/starch solutions are applied to the paper web by means of various mechanical presses (size presses). Together with surface sizing agents the surface starches impart additional strength to the paper web and additionally provide water hold out or "size" for superior printing properties. Starch is also used in paper coatings as one of the binders for the coating formulations which include a mixture of pigments, binders and thickeners. Coated paper has improved smoothness, hardness, whiteness and gloss and thus improves printing characteristics.

Corrugated board adhesives

Corrugated board adhesives are the next largest application of non-food starches globally. Starch glues are mostly based on unmodified native starches, plus some additive such as borax and caustic soda. Part of the starch is gelatinized to carry the slurry of uncooked starches and prevent sedimentation. This opaque glue is called a SteinHall adhesive. The glue is applied on tips of the fluting. The fluted paper is pressed to paper called liner. This is then dried under high heat, which causes the rest of the uncooked starch in glue to swell/gelatinize. This gelatinizing makes the glue a fast and strong adhesive for corrugated board production.

Clothing starch

Clothing or laundry starch is a liquid prepared by mixing a vegetable starch in water (earlier preparations also had to be boiled), and is used in the laundering of clothes. Starch was widely used in Europe in the 16th and 17th centuries to stiffen the wide collars and ruffs of fine linen which surrounded the necks of the well-to-do. During the 19th and early 20th century it was stylish to stiffen the collars and sleeves of men's shirts and the ruffles of women's petticoats by applying starch to them as the clean clothes were being ironed. Starch gave clothing smooth, crisp edges, and had an additional practical purpose: dirt and sweat from a person's neck and wrists would stick to the starch rather than to the fibers of the clothing. The dirt would wash away along with the starch; after laundering, the starch would be reapplied. Today, starch is sold in aerosol cans for home use.

Other

Another large non-food starch application is in the construction industry, where starch is used in the gypsum wall board manufacturing process. Chemically modified or unmodified starches are added to the stucco containing primarily gypsum. Top and bottom heavyweight sheets of paper are applied to the formulation, and the process is allowed to heat and cure to form the eventual rigid wall board. The starches act as a glue for the cured gypsum rock with the paper covering, and also provide rigidity to the board.

Starch is used in the manufacture of various adhesives or glues for book-binding, wallpaper adhesives, paper sack production, tube winding, gummed paper, envelope adhesives, school glues and bottle labeling. Starch derivatives, such as yellow dextrins, can be modified by addition of some chemicals to form a hard glue for paper work; some of those forms use borax or soda ash, which are mixed with the starch solution at 50–70 °C (122–158 °F) to create a very good adhesive. Sodium silicate can be added to reinforce these formula.

• Textile chemicals from starch: warp sizing agents are used to reduce breaking of yarns during weaving. Starch is mainly used to size cotton based yarns. Modified starch is also used as textile printing thickener.
• In oil exploration, starch is used to adjust the viscosity of drilling fluid, which is used to lubricate the drill head and suspend the grinding residue in petroleum extraction.
• Starch is also used to make some packing peanuts, and some drop ceiling tiles.
• In the printing industry, food grade starch is used in the manufacture of anti-set-off spray powder used to separate printed sheets of paper to avoid wet ink being set off.
• For body powder, powdered corn starch is used as a substitute for talcum powder, and similarly in other health and beauty products.
• Starch is used to produce various bioplastics, synthetic polymers that are biodegradable. An example is polylactic acid based on glucose from starch.
• Glucose from starch can be further fermented to biofuel corn ethanol using the so-called wet milling process. Today most bioethanol production plants use the dry milling process to ferment corn or other feedstock directly to ethanol.
• Hydrogen production could use glucose from starch as the raw material, using enzymes.

Occupational safety and health

The Occupational Safety and Health Administration (OSHA) has set the legal limit (Permissible exposure limit) for starch exposure in the workplace as 15 mg/m³ total exposure and 5 mg/m³ respiratory exposure over an 8-hour workday. The National Institute for Occupational Safety and Health (NIOSH) has set a Recommended exposure limit (REL) of 10 mg/m³ total exposure and 5 mg/m³ respiratory exposure over an 8-hour workday.