Etymology[edit]

In Mexico, corn smut is known as huitlacoche (Spanish pronunciation: [(ɡ)witɬaˈkotʃe], sometimes

spelled cuitlacoche). This word entered Spanish in Mexico from Classical Nahuatl, though the
Nahuatl words from which huitlacoche is derived are debated. In modern Nahuatl, the word
for huitlacoche is cuitlacochin (Nahuatl pronunciation: [kʷit͡ɬɑˈkot͡ʃin]), and some sources
deem cuitlacochi to be the classical form.[5]
Some sources wrongly give the etymology as coming from the Nahuatl
words cuitlatl [ˈkʷit͡ɬɑt͡ɬ] ("excrement" or "rear-end", actually meaning "excrescence")
and cochtli [ˈkot͡ʃt͡ɬi] ("sleeping", from cochi "to sleep"), thus giving a combined mismeaning of
"sleeping/hibernating excrement",[5][6] but actually meaning "sleeping excrescence", referring to the
fact that the fungus grows in between the corns and impedes them from developing, thus they
remain "sleeping".
A second group of sources deem the word to mean "raven's excrement".[7][8] These sources appear to
be combining the word cuitlacoche for "thrasher"[9] with cuitla, meaning "excrement", actually
meaning "excrescence". However, the avian meaning of cuitlacoche derives from the Nahuatl word
"song" cuīcatl [ˈkʷiːkɑt͡ɬ], itself from the verb "to sing" cuīca [ˈkʷiːkɑ].[5] This root then clashes with this
reconstruction's second claim that the segment cuitla- comes from cuitla ("excrement").
One source derives the meaning as "corn excrescence", using cuītla again and "maize" tlaōlli [t͡ɬɑ
ˈoːlːi].[10] This requires the linguistically unlikely evolution of tlaole "maize" into tlacoche.

Taxonomy[edit]
U. maydis is the best known and studied of the Ustilaginomycetes and so is often used as the
exemplar species when talking about its entire class.[11]

Characteristics[edit]
The fungus infects all parts of the host plant by invading the ovaries of its host. The infection causes
the corn kernels to swell up into tumor-like galls, whose tissues, texture, and developmental pattern
are mushroom-like. These galls are made up of hypertrophied cells of the infected plant, along with
resulting fungal threads, and blue-black spores.[12] These dark-colored spores give the cob a burned,
scorched appearance; this is the origin of the generic name Ustilago, from the Latin word ustilare (to
burn).
This ear of corn has been infected with ustilago maydis.

Biology[edit]
Life cycle[edit]

Ustilago maydis haploid sporidia

When grown in the lab on very simple media, it behaves like baker's yeast, forming
single cells called sporidia. These cells multiply by budding off daughter cells. When two compatible
sporidia meet on the surface of the plant, they switch to a different mode of growth. First, they
produce one or another pheromone, and begin producing one or the other type of pheromone
receptor - this depends on mating type a or b, as determined by alleles at two unlinked mating loci. If
this signalling is successful they then send out conjugation tubes to find each other,[11] after which
they fuse and make a hypha to enter the maize plant. Hyphae growing in the plant are dikaryotic;
they possess two haploid nuclei per hyphal compartment. In contrast to sporidia, the dikaryotic
phase of U. maydis requires infection of the plant to grow and differentiate, and cannot be
maintained in the laboratory.
Proliferation of the fungus inside the plant leads to disease symptoms such
as chlorosis, anthocyanin formation, reduced growth, and the appearance of tumors harboring the
developing teliospores.[13][14]
Mature tumors release spores that rain and wind then disperse. Under appropriate conditions,
a metabasidium is formed in which meiosis occurs. Resulting haploid nuclei migrate into elongated
single cells. These cells detach from the metabasidium to become the sporidia, thus completing the
life cycle.
Host/pathogen conflict[edit]
Plants have evolved efficient defense systems against pathogenic microbes. A rapid plant defense
reaction after pathogen attack is the oxidative burst, which involves the production of reactive
oxygen species at the site of the attempted invasion. As a pathogen, U. maydis can respond to such
an oxidative burst by an oxidative stress response, regulated by gene YAP1. This response
protects U. maydis from the host attack, and is necessary for the pathogen’s virulence.
[15]
 Furthermore, U. maydis has a well-established recombinational DNA repair system.[16] This repair
system involves a homolog of Rad51 that has a very similar sequence and size to its mammalian
counterparts. This system also involves a protein, Rec2 that is more distantly related to Rad51, and
Brh2 protein that is a streamlined version of the mammalian Breast Cancer 2 (BRCA2) protein.
When any of these proteins is inactivated, sensitivity of U. maydis to DNA damaging agents is
increased. Also mitotic recombination becomes deficient, mutation frequency increases and meiosis
fails to complete. These observations suggest that recombinational repair during mitosis and meiosis
in U. maydis may assist the pathogen in surviving DNA damage arising from the host’s oxidative
defensive response to infection, as well as from other DNA damaging agents.

Proteome[edit]
U. maydis is known to produce four Gα proteins, and one each of Gβ and Gγ.[11]

Management[edit]
There are many ways to control and manage corn smut; however, corn smut cannot be controlled by
any common fungicide at this time, as Ustilago maydis infects individual corn kernels instead of
infecting the entire cob, like head smut.[17] Some beneficial ways to contain corn smut include
resistant corn plants, crop rotation, and avoiding mechanical injury to the plant. A mechanical injury
can cause the corn to become easily accessible to Ustilago maydis, enhancing infection.
Additionally, clearing the planting area of debris can help control corn smut, as the teliospores from
corn smut overwinter in debris. This is not the best practice, though, because corn smut can also
overwinter in the soil; crop rotation is recommended. Lastly, as excess nitrogen in the soil augments
infection rate, using fertilizer with low nitrogen levels, or just limiting the amount of nitrogen in the soil
proves to be another way to control corn smut.[18]

Ear of corn infected with Ustilago maydis

Environment[edit]
Although not all the conditions that favor growth of Ustilago maydis are known, there are certain
environments where corn smut seems to thrive, depending on both abiotic and biotic factors. Hot
and dry weather during pollination followed by a heavy rainy season appear to improve the
pathogenicity of corn smut.[19] Furthermore, excess manure (and therefore nitrogen) in the soil also
increases pathogenicity. Not only do these abiotic factors increase infectability, they also increase
disease spread. High winds and heavy rain also increase disease spread as the spores of corn smut
can be more easily transmitted. Other biotic factors largely have to do with the extent by which
humans interact with the corn and corn smut. If corn debris is not cleared at the end of the season,
the spores can overwinter in the corn fragments and live to infect another generation.[20] Finally,
humans wounding the corn (with shears or other tools of the like) present the opportunity for corn
smut to easily enter the plant.

Uses[edit]
Model organism[edit]
The yeast-like growth of U. maydis makes it an appealing model organism for research, although its
relevance in nature is unknown. The fungus is exceptionally well-suited for genetic modification. This
allows researchers to study the interaction between the fungus and its host with relative ease. The
availability of the entire genome is another advantage of this fungus as model organism.[21]
U. maydis is not only used to study plant disease, but it also is used to study plant genetics. In 1996,
a study on U. maydis genetics led to the discovery of synthesis-dependent strand annealing, a
method of homologous recombination used in DNA repair.[22] Other studies in the fungus have also
investigated the role of the cytoskeleton in polarized growth.[citation needed] It is largely due to work with U.
maydis that the function of the breast-cancer gene BRCA2 is now known.[23] The fungus is mostly
studied as model organism for host pathogen interaction and delivery of effectors protein

Industrial biotechnology[edit]
Ustilago maydis is able to produce a broad range of valuable chemicals such as ustilagic
acid, itaconic acid, malic acid, and hydroxyparaconic acid. With this ability it is gaining more and
more relevance for industrial applications.[24]

Culinary uses[edit]
See also: List of delicacies
Smut feeds on the corn plant and decreases the yield. Smut-infected crops are often destroyed,
although some farmers use them to prepare silage. However, the infected galls are still edible, and
in Mexico they are highly esteemed as a delicacy, where it is known as huitlacoche, being preserved
and sold for a significantly higher price than uninfected corn. The consumption of corn smut
originated directly from Aztec cuisine.[25] For culinary use, the galls are harvested while still immature
— fully mature galls are dry and almost entirely spore-filled. The immature galls, gathered two to
three weeks after an ear of corn is infected, still retain moisture and, when cooked, have a flavor
described as mushroom-like, sweet, savory, woody, and earthy. Flavor compounds
include sotolon and vanillin, as well as the sugar glucose.
Huitlacoche is a source of the essential amino acid lysine, which the body requires but cannot
manufacture. It also contains levels of beta-glucans similar to, and protein content equal or superior
to, most edible fungi.[26]
The fungus has had difficulty entering into the American and European diets as most farmers see it
as blight, despite attempts by government and high-profile chefs to introduce it. In the mid-1990s,
due to demand created by high-end restaurants, Pennsylvania and Florida farms were allowed by
the United States Department of Agriculture (USDA) to intentionally infect corn with huitlacoche.
Most observers consider the program to have had little impact,[citation needed] although the initiative is still in
progress. The cursory show of interest is significant because the USDA has spent a considerable
amount of time and money trying to eradicate corn smut in the United States. Moreover, in 1989,
the James Beard Foundation held a high-profile huitlacoche dinner, prepared by Josefina Howard,
chef at Rosa Mexicano restaurant.[27] This dinner tried to get Americans to eat more of it by renaming
it the Mexican truffle and it is often compared to truffles in food articles describing its taste and
texture.[27][28][29]
Native Americans of the American Southwest, including the Zuni people, have used corn smut in an
attempt to induce labor. It has similar medicinal effects to ergot, but weaker, due to the presence of
the chemical ustilagine.[30]

Huitlacoche corn taco
 

Quesadilla de huitlacoche, as it is often served in central Mexico

 

Huitlacoche for sale in the produce department of a Soriana store in Mexico

Recipes of Mexico[edit]
A simple Mexican-style succotash can be made from chorizo, onions, garlic, serrano
peppers, huitlacoche, and shrimp with salsa taquera. The mild, earthy flavors of
the huitlacoche blend nicely with the fats of the chorizo and bond to mellow out the heat from the
peppers and salsa.
Another Mayan favorite on the Riviera Maya (Cancun to Tulum) is to add huitlacoche to omelettes.
Once again, its earthy flavors bond with the fats that cook the eggs to mellow the flavors into a
truffle-like taste.
Huitlacoche is also popular in quesadillas with Mexican cheese, sautéed onions, and tomatoes.
The blueish color transforms into the recognizable black color only with heat. Any dish
with huitlacoche must include a slow simmer of the fungus until it becomes black, which also
removes most of the starch of the corn, and what is left is a black oily paste.
Availability[edit]
In Mexico, huitlacoche is mostly consumed fresh and can be purchased at restaurants or street or
farmer's markets throughout the country and, to a much lesser extent, can also be purchased as a
canned good in some markets and via the internet. Farmers in the countryside spread the spores
around intentionally to create more of the fungus. In some parts of the country, they call the fungus
"hongo de maiz", i.e. "maize fungus".[31]