Blakeslea Trispora
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''Blakeslea trispora'' is a
mould A mold () or mould () is one of the structures that certain fungi can form. The dust-like, colored appearance of molds is due to the formation of spores containing fungal secondary metabolites. The spores are the dispersal units of the fungi ...
and member of the division
Zygomycota Zygomycota, or zygote fungi, is a former phylum, division or phylum of the kingdom Fungi. The members are now part of two Phylum, phyla: the Mucoromycota and Zoopagomycotina, Zoopagomycota. Approximately 1060 species are known. They are mostly t ...
. This species has been well studied for its ability to produce carotenoids, particularly, β-carotene and
lycopene Lycopene is an organic compound classified as a tetraterpene and a carotene. Lycopene (from the Neo-Latin '' Lycopersicon'', the name of a former tomato genus) is a bright red carotenoid hydrocarbon found in tomatoes and other red fruits and ve ...
. β-carotene is a vitamin A precursor and both of β-carotene and lycopene play a significant role in the inhibition of
oxidative stress Oxidative stress reflects an imbalance between the systemic manifestation of reactive oxygen species and a biological system's ability to readily detoxify the reactive intermediates or to repair the resulting damage. Disturbances in the normal ...
. ''Blakeslea trispora'' is commonly isolated from soil samples throughout the Southern United States and Southern Asia. ''B. trispora'' is a
pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
of tropical plants.
In vivo Studies that are ''in vivo'' (Latin for "within the living"; often not italicized in English) are those in which the effects of various biological entities are tested on whole, living organisms or cells, usually animals, including humans, an ...
pathogenicity testing using
animal models A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mod ...
suggests this fungus is not a cause of animal or human disease.


History

The genus ''Blakeslea'' was named in honor of American botanist Albert Francis Blakeslee based on ''B. trispora''. While A.F. Blakeslee was studying another fungus called
Botrytis rileyi
', he collaborated with
Roland Thaxter Roland Thaxter (August 28, 1858 – April 22, 1932) was an American mycologist, plant pathologist, botanist, and entomologist, renowned for his contribution to the insect parasitic fungi—Laboulbeniales. His college education was completed at Har ...
to further study this fungus. In 1914, while Thaxter was doing further research on ''Botrytis rileyi,'' ''Blakeslea trispora'' was accidentally isolated from a contaminated caterpillar infected with ''Botrytis rileyi''. ''Blakeslea trispora'' was first identified from the
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e of the caterpillar, which was growing on the
cowpea The cowpea (''Vigna unguiculata'') is an annual herbaceous legume from the genus '' Vigna''. Its tolerance for sandy soil and low rainfall have made it an important crop in the semiarid regions across Africa and Asia. It requires very few inpu ...
plant. The caterpillar was infected by the fungus ''Botrytis rileyi;'' however, ''Blakeslea trispora'' was thought to be incidentally transferred to the diseased caterpillar feeding on a cowpea flower. When Thaxter first identified ''B. trispora'', he considered ''B. trispora'' to be very closely related to the genus '' Choanephora'' because of highly similar sporangiospore morphology. Both have a distinctive brown colour with faint, longitudinal striations on the sporangiole wall. The shape of the large spherical heads of their sporangiola are also similar. However, ''Choanephora'' and ''Blakeslea'' are considered to be distinct genera and can be distinguishable by the spore wall and its separation from the sporangiole wall. Species of ''Choanephora'' have highly adherent sporangiole wall in contrast to ''Blakeslea'' species where the sporangiole wall is readily separable from the underlying spore at maturity.


Growth and morphology

''Blakeslea trispora'' undergoes both sexual and asexual reproduction. The asexual reproductive phase of ''Blakeslea trispora'' involves the production of sporangiospores produced in
sporangia A sporangium (from Late Latin, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...
. Once released, they can germinate in the presence of free water. Colonies of ''B. trispora'' grow rapidly on the agar growth media at 25 Â°C. They are white at first but become yellow to pale brown and very dark brown as they mature. The hyphae of ''B. trispora'' are aseptate, very dense, and highly branched. Sexual reproduction is by the formation of
zygospore A zygospore is a diploid reproductive stage in the life cycle of many fungi and protists. Zygospores are created by the nuclear fusion of haploid cells. In fungi, zygospores are formed in zygosporangia after the fusion of specialized budding stru ...
s, which contain high concentrations of triglycerol-rich
lipid Lipids are a broad group of organic compounds which include fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include storing ...
s and
phosphatidylcholine Phosphatidylcholines (PC) are a class of phospholipids that incorporate choline as a headgroup. They are a major component of biological membranes and can easily be obtained from a variety of readily available sources, such as egg yolk or soyb ...
. Zygospores can persist for long periods of time, and their germination is dependent on a cytoplasmic regulatory system that sustains dormancy and forestalls germination in the presence of unfavorable growth conditions. Zygospores range in size from 40-80μm. They are spherical or slightly flattened in shape. ''Blakeslea trispora'' has a
heterothallic Heterothallic species have sexes that reside in different individuals. The term is applied particularly to distinguish heterothallic fungi, which require two compatible partners to produce sexual spores, from homothallic ones, which are capable ...
mating system, having (+) and (-) mating types. Contact and interchange between the opposite mating types is a necessary precursor to induce sexual reproduction and development of zygospores. Extensions called
gametangia A gametangium (: gametangia) is a sex organ or cell in which gametes are produced that is found in many multicellular protist A protist ( ) or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists ...
are formed from each of the compatible
haploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
mycelia. Following anastomosis, a fertile
heterokaryotic In biology, a heterokaryon is a multinucleate cell that contains genetically different nuclei. This is a special type of syncytium. This can occur naturally, such as in the mycelium of fungi during sexual reproduction, or artificially as formed by ...
zygosporangium A zygospore is a diploid reproductive stage in the life cycle of many fungi and protists. Zygospores are created by the nuclear fusion of haploid cells. In fungi, zygospores are formed in zygosporangia after the fusion of specialized budding stru ...
is formed within which the zygospores develop. During sexual reproduction, carotenoid pigments are produced by both of mating type.
Carotenoid Carotenoids () are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, archaea, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, corn, tomatoes, cana ...
s are precursors of many
apocarotenoid Apocarotenoids are organic compounds which occur widely in living organisms. They are derived from carotenoids by oxidative cleavage, catalyzed by carotenoid oxygenases. Examples include the vitamin A retinoids retinal, retinoic acid, and retinol; a ...
s that contain very important sex-specific precursors, trisporic acid (TSA) for the sexual reproduction of ''Blakeslea trispora''.
Carotene The term carotene (also carotin, from the Latin ''carota'', "carrot") is used for many related unsaturated hydrocarbon substances having the formula C40Hx, which are synthesized by plants but in general cannot be made by animals (with the ex ...
s produced from carotenoids are further processed by carotene oxygenase to synthesize trisporic acid (TSA). TSA produced from carotene stimulates both sexually complementary cells to make contact with each other. TSA is considered an important
signalling molecule In biology, cell signaling (cell signalling in British English) is the process by which a cell interacts with itself, other cells, and the environment. Cell signaling is a fundamental property of all cellular life in both prokaryotes and eukaryot ...
for the initiation and control sexual reproduction.


Physiology

At the beginning of the sexual reproduction cycle of ''B. trispora,'' the initial step is the production of carotenes from carotenoids. Carotenes are further processed by carotene oxygenase, which is encoded in the tsp3 gene of the ''B. trispora'', to produce TSA. TSA is produced by both of the mating types: (+) and (-) strains, and it is copiously produced especially when compatible mycelia are grown together. As these two different sex types produce TSA, they sense sexually complementary cells and form gametangia. Eventually, those gametangia merge and zygosporangia form. As these two different mating types meet each other, each mating type transfers the sex-specific precursor of the trisporoid TSA, and acts as a signal for the synthesis of the surface protein
agglutinin An agglutinin is a substance in the blood that causes particles to coagulate and aggregate; that is, to change from fluid-like state to a thickened-mass (solid) state. Agglutinins can be antibodies that cause antigens to aggregate by binding t ...
. Agglutinin allows the two TSAs to recognize each other. It then causes rapid contact and efficient interactions between those two different mating types. Additionally, stimulating both of the mating types by TSA promotes synthesis of β-carotene. As β-carotene is produced, it becomes a precursor of trisporoid, which is a pheromone for ''B. trispora''. Production of β-carotene promotes a
positive feedback Positive feedback (exacerbating feedback, self-reinforcing feedback) is a process that occurs in a feedback loop where the outcome of a process reinforces the inciting process to build momentum. As such, these forces can exacerbate the effects ...
process that further stimulates carotenogenesis and the production of trisporoid which serves as a β-carotene increasing substance. Furthermore, it act as a hormone stimulator of its
biosynthesis Biosynthesis, i.e., chemical synthesis occurring in biological contexts, is a term most often referring to multi-step, enzyme-Catalysis, catalyzed processes where chemical substances absorbed as nutrients (or previously converted through biosynthe ...
. Thus, ''Blakeslea trispora'' requires certain concentrations of TSA to activate carotenogenesis and produce more carotenoids (about 0.5% of its
dry weight Vehicle weight is a measurement of wheeled motor vehicles; either an actual measured weight of the vehicle under defined conditions or a gross weight rating for its weight carrying capacity. Curb or kerb weight Curb weight (American English) or k ...
) which can be accumulated in the zygospores of ''B. trispora''. Therefore, both TSA and trisporoid acts as sex hormones in ''Blakeslea trispora'', which triggers the sexual reproduction and controls intimate contact between heterothallic strains, further governing the formation of sex structures, zygospores. Carotenoids are absolutely necessary not only for the production of trisporic acid, but also for the process of zygote formation, as significant factors for the production of sporopollenin, a structural component of the zygospore cell wall. It is consequential to regulate this feedback-type synthesis with carotenoids and further synthesis of TSA. Therefore, the formation of the zygospores can be prevented by the inhibition of carotenogenesis in ''Blakeslea trispora''.


Applications

''Blakeslea trispora'' is useful as sources of β-carotene and its precursor molecule, lycopene, in industrial production. These molecules are useful
food coloring Food coloring, color additive or colorant is any dye, pigment, or substance that imparts color when it is added to food or beverages. Colorants can be supplied as liquids, powders, gels, or pastes. Food coloring is commonly used in commercia ...
agents and may have beneficial effects for human health as
antioxidant Antioxidants are Chemical compound, compounds that inhibit Redox, oxidation, a chemical reaction that can produce Radical (chemistry), free radicals. Autoxidation leads to degradation of organic compounds, including living matter. Antioxidants ...
s. The production of lycopene primarily requires some interaction between the mating strains. ''Blakeslea trispora'' needs both mating types to synthesize lycopene on a commercially applicable scale. The (-) strain is twice proportionally as important as the (+) strain in determining the productivity of the synthesis of the lycopene. To produce an optimal amount of lycopene, excess (-) mating type at a 1:2 (+/-) ratio with inoculum ages 36 and 48 hours respectively is favorable.


Lycopene

''Blakeslea trispora'' is known to be the most effective producer of lycopene. Lycopene is processed by lycopene cyclase which leads to the production of β-carotene. For industrial production of lycopene, ''Blakeslea trispora'' is grown with an lycopene cyclase inhibitor which can be introduced into the
fermentation Fermentation is a type of anaerobic metabolism which harnesses the redox potential of the reactants to make adenosine triphosphate (ATP) and organic end products. Organic molecules, such as glucose or other sugars, are catabolized and reduce ...
process. The zygospores of ''Blakeslea trispora'' tend to contain a maximum amount of lycopene. Lycopene is an intermediate in the biosynthesis of all dicyclic carotenoids including β-carotene. Lycopene is one of the most important carotene molecules because it is capable of producing both β-carotene and other carotenoids, well known for their potent
anti-oxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. Autoxidation leads to degradation of organic compounds, including living matter. Antioxidants are frequently added to industrial products, s ...
activities. As such, β-carotene and other carotenoids play crucial roles for
oxidative stress Oxidative stress reflects an imbalance between the systemic manifestation of reactive oxygen species and a biological system's ability to readily detoxify the reactive intermediates or to repair the resulting damage. Disturbances in the normal ...
reduction and cardiovascular protection. Carotenoids have highly efficient antioxidant scavenging activities against ROS (
reactive oxygen species In chemistry and biology, reactive oxygen species (ROS) are highly Reactivity (chemistry), reactive chemicals formed from diatomic oxygen (), water, and hydrogen peroxide. Some prominent ROS are hydroperoxide (H2O2), superoxide (O2−), hydroxyl ...
), such as singlet-oxygen and
free radicals In chemistry, a radical, also known as a free radical, is an atom, molecule, or ion that has at least one unpaired electron, unpaired valence electron. With some exceptions, these unpaired electrons make radicals highly chemical reaction, chemi ...
. Therefore, they have the ability to prevent
chronic diseases A chronic condition (also known as chronic disease or chronic illness) is a health condition or disease that is persistent or otherwise long-lasting in its effects or a disease that comes with time. The term ''chronic'' is often applied when the ...
such as cancer, cerebrovascular and cardiovascular diseases and
myocardial infarction A myocardial infarction (MI), commonly known as a heart attack, occurs when Ischemia, blood flow decreases or stops in one of the coronary arteries of the heart, causing infarction (tissue death) to the heart muscle. The most common symptom ...
. Lycopene is considered a very important and relevant source to human health. A case study by Weilian Hu and his colleagues in 2013 showed that the administration of lycopene in adult mice appeared to improve the activity of antioxidant enzyme. They have reported that, the administration of ''Blakeslea trispora'' powder, which contains high amounts of lycopene has the potential to protect the liver, brain, kidney and skin against oxidative stress. This is done by reducing the concentration of ROS and by enhancing the activities of the antioxidant enzyme. Furthermore, they are further investigating whether the fungus ''Blakeslea trispora'' could be a potent effector of anti-aging because of its ability to efficiently mass-produce amounts of lycopene.


β-carotene

β-carotene is a molecule which displays a red-orange pigment. Therefore, it is used as a coloring agent for food products. β-carotene is a member of carotenes which are highly unsaturated
isoprene Isoprene, or 2-methyl-1,3-butadiene, is a common volatile organic compound with the formula CH2=C(CH3)−CH=CH2. In its pure form it is a colorless volatile liquid. It is produced by many plants and animals (including humans) and its polymers ar ...
derivatives. Because ''Blakeslea trispora'' has an effective ability to produce great amount of β-carotene from lycopene, ''Blakeslea trispora'' is the main organism used for its production on an industrial scale. β-carotenes are known to be a powerful stimulant of the human
immune system The immune system is a network of biological systems that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to bacteria, as well as Tumor immunology, cancer cells, Parasitic worm, parasitic ...
and play significant roles in the prevention of
degenerative disease Degenerative disease is the result of a continuous process based on degenerative cell changes, affecting tissues or organs, which will increasingly deteriorate over time. In neurodegenerative diseases, cells of the central nervous system stop wo ...
s and cancers. All cells are capable of producing and regulating ROS. However, dysregulation of ROS can lead to
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
damaging, inactivation of enzymes and proteins, disruption of membranes. This ultimately causes cell death, becoming very toxic to the individuals. Further investigations of β-carotene usage collected from ''Blakeslea trispora'' may lead to great improvements to human health in the treatment and prevention of certain chronic diseases such as cancer.


References

{{DEFAULTSORT:Blakeslea Trispora Fungi described in 1914 Fungal plant pathogens and diseases Zygomycota Fungus species