Crucibulum
Encyclopedia
Crucibulum is a genus
in the Nidulariaceae
, a family
of fungi whose fruiting bodies resemble tiny egg-filled bird's nests. Often called "splash cups", the fruiting bodies are adapted for spore
dispersal by using the kinetic energy
of falling drops of rain. The "eggs" inside the bird's nests (technically known as peridioles) are hard waxy shells containing spores, and tend to stick to whatever nearby herbage they land on, thus increasing the odds of being consumed and dispersed by herbivorous animals. Members of this genus are saprobic, obtaining nutrients from dead organic matter
, and are typically found growing on decayed wood and wood debris. The three known Crucibulum species (C. laeve, C. parvulum, and C. cyathiforme) are distinguished from other genera of the Nidulariaceae by their relatively simple funiculus – a cord of hyphae that connects the peridiole (the "eggs") to the exterior of the bird's nest.
The structure and biology of the genus Crucibulum was better known by the mid-19th century, when the brothers Louis René
and Charles Tulasne
published a monograph
on the bird's nest fungi. Subsequently, monographs were written in 1902 by Violet S. White (American species), Curtis Gates Lloyd
in 1906, Gordon Herriot Cunningham
in 1924 (New Zealand species), and Harold J. Brodie
in 1975.
The type species
for the genus Crucibulum described by the Tulasne brothers was Crucibulum vulgare, an older synonym of the species known today as C. laeve. However, this naming choice was later deemed invalid by rules of fungal nomenclature; the first name validly applied to the species was C. laeve, use by De Candolle, who had based his species upon Nidularia laevis as it appeared in Bulliard's
Histoire des Champignons de la France (Paris, 1791). Kambly and Lee published the first taxonomically
valid description of the genus in 1936. In their 1844 monograph on the Nidulariaceae, the brothers Louis René
and Charles Tulasne
used the name Crucibulum vulgare, and the species was known by this name until the International Commission on the Taxonomy of Fungi (ICTF) changed the starting-point date for the naming of fungi, and C. vulgare was deemed invalid. The etymology
of the specific epithet is derived from the Latin laeve, meaning "smooth".
to cinnamon-colored fruiting bodies
, known as a peridium
, that are cup- or crucible
-shaped. Depending on the species, the size of the peridium may range from 2–4 tall by 1.5–3 mm wide at the mouth (for C. parvulum) to 5–10 mm tall by 5–8 mm wide (for C. laeve). Viewed microscopically, the wall of the peridium is made of a single layer of tissue, in contrast to the three-layered peridium wall in Cyathus species. The outer surface of the peridium has hypha
e that agglutinate so as to form a texture with visible filaments, a condition known as fibrillose; this outer layers of hairs typically wears off with age to leave a relatively smooth surface.
Young specimens have a thin layer of tissue called an epiphragm that covers the top of the peridium; it wears off at maturity to expose the peridioles within. There are usually 4–6 peridioles (up to 15 have been noted for C. laeve) that are disc-shaped, whitish in color, and attached to the endoperidium by a strand called a funicular cord. Made of mycelia
, The funicular cord tends to wither away and disappear as the fruiting body ages. Spore
s from Crucibulum species typically have an elliptical or roughly spherical shape, and are thick-walled, transclucent (hyaline
) or light yellow-brown in color, with dimensions of 5–15 by 5–8 µm
. the spores of C. cyathiforme are notably slightly or strongly curved.
Because the basic fruiting body structure in all genera
of the Nidulariaceae family is essentially similar, Crucibulum may be readily confused with species of Nidula
or Cyathus
, especially older, weathered specimens of Cyathus that may have the hairy ectoperidium worn off. It distinguished from Nidula by the presence of a funiculus, a cord of hyphae attaching the peridiole to the endoperidium. Cyathus differs from genus Crucibulum by having a distinct three-layered wall and a more intricate funiculus.
word peridion, meaning "small leather pouch", the peridiole is the "egg" of the bird's nest. It is a mass of basidiospores and gleba
l tissue enclosed by a hard and waxy outer shell. In Crucibulum, the disc-shaped peridioles are light buff or white; the is due to a persistent layer of tissue surrounding the peridioles, called a tunica. Inside the peridiole is a spore-bearing tissue (the hymenium
) that is made of spore-bearing cells (basidia), sterile (non-reproductive) structures, and spores.
Peridioles are attached to the fruiting body by a funiculus, a structure of hypha
e that may be differentiated into three regions: the basal piece, which attaches it to the inner wall of the peridium, the middle piece, and an upper sheath, called the purse, connected to the lower surface of the peridiole. In the purse and middle piece is a coiled thread of interwoven hyphae called the funicular cord, attached at one end to the peridiole and at the other end to an entangled mass of hyphae called the hapteron. In Crucibulum species the peridioles is covered by a whitish tunica. The funiculus of Crucibulum species is markedly different from those of Cyathus species: in Crucibulum, the purse is a rounded knob 0.3–0.5 mm wide, attached to the underside of the peridiole. Attaching the purse directly to the wall of the peridium is a stout yellow-grey cord 0.1 mm wide and about 2.5 mm long.
Spores are dispersed when a peridiole is dislodged by raindrops or water dripping off an over-hanging leaf. The smooth inner walls of the fruiting body consistently form an angle of 70–75° with the horizontal; it has been demonstrated experimentally that the combined effect of the crucible shape and internal wall angle produce a good splash action. The force of the falling water splashes out the peridiole, uncoiling and snapping the funiculus, the cord that connects it to the fruiting body. As the peridiole continues its flight, the funiculus extends to its full length. The sticky end of the funiculus may adhere to a leaf or a twig some distance away, and the peridiole may end up being wrapped around or hanging down the object to which the funiculus is stuck. The spores can germinate when the thick outer wall of the peridiole wears away, or the peridiole may be eaten by a herbivorous animal, and ultimately passed through its digestive system. This method of spore dispersal, first suggested by John Ray
in the late 17th century, was tested experimentally by Martin (1927), and more thoroughly by Buller
and Brodie
in the 1940s.
that can reproduce both asexually (via vegetative
spores), or sexually (with meiosis
). Like other wood-decay fungi, this life cycle may be considered as two functionally different phases: the vegetative stage for the spread of mycelia, and the reproductive stage for the establishment of spore-producing structures, the fruiting bodies.
The vegetative stage encompasses those phases of the life cycle involved with the germination, spread, and survival of the mycelium. Spores germinate under suitable conditions of moisture and temperature, and grow into branching filaments called hypha
e, pushing out like roots into the rotting wood. These hyphae are homokaryotic
, containing a single nucleus in each compartment; they increase in length by adding cell-wall material to a growing tip. As these tips expand and spread to produce new growing points, a network called the mycelium develops. Mycelial growth occurs by mitosis
and the synthesis of hyphal biomass. When two homokaryotic hyphae of different mating compatibility groups
fuse with one another, they form a dikaryotic
mycelia in a process called plasmogamy
. Prerequisites for mycelial survival and colonization a substrate (like rotting wood) include suitable humidity and nutrient availability.Crucibulum laeve is saprobic, so mycelial growth in rotting wood is made possible by the secretion of enzyme
s that break down complex polysaccharide
s (such as cellulose
and lignin
) into simple sugars that can be used as nutrients.
After a period of time and under the appropriate environmental conditions, the dikaryotic mycelia may enter the reproductive stage of the life cycle. Fruiting body formation is influenced by external factors such as season (which affects temperature and air humidity), nutrients and light. As fruiting bodies develop they produce peridioles containing the basidia upon which new basidiospores are made. Young basidia contain a pair of haploid sexually compatible nuclei which fuse, and the resulting diploid fusion nucleus undergoes meiosis to produce basidiospore
s, each containing a single haploid nucleus. The dikaryotic mycelia from which the fruiting bodies are produced is long lasting, and will continue to produce successive generations of fruiting bodies as long as the environmental conditions are favorable.
s are produced on club-shaped basidia which line the internal cavity of the peridiole. Basidia typically have 4 spores, attached by a short projection (a sterigma
); after being detached from the basidia the spores migrate towards the center of the peridiole concurrently with the collapse and gelatinization of the underlying tissues.
Grown in liquid culture, Crucibulum laeve produces bioactive chemicals called salfredins that are structurally
related to benzofuran
and chromene, molecules that contain cyclic amide
or lactone
five-membered ring structures; these compounds are unique to this species. Salfredin B11 was first identified in 1995, while later research confirmed the presence of additional salfredin-type metabolites
. These compounds are inhibitors
of aldose reductase
, an enzyme
that has been implicated in the formation of cataract
s in advanced stages of diabetes mellitus
. The salfredin compounds may have therapeutic use in the treatment of this disorder.
. They are typically found growing on wood and woody debris such as stems, twigs, wood chips, old nut shells, and old matting; they are sometimes found on "dried manure cakes". Brodie notes (of C. laeve) they are "never" found on soil or large logs. C. parvulum has been found on the roots and stems of old or dead dry land plants such as Juniperus horizontalis
and Artemisa
species.
C. laeve, the most well-known species of Crucibulum, is a temperate
-zone species with a circumpolar distribution. It has been collected in most European countries and the Canary Islands
; in North America it has been found from Alaska to Mexico, while South American locations include Chile and Tierra del Fuego
. It has also been found in Australia, Iceland, Japan, and New Zealand. C. parvulum has a primarily North American distribution, having been found in Alaska
, southern Alberta
Badlands
, the Canadian Rocky Mountains
and in semi-deserts of Idaho
; in 2004 it was collected in China. C. cyathiforme is only known from Colombia—where it was discovered growing at an elevation of nearly 7000 feet (2146 metres)— and Armenia
.
s or other substances considered toxic to humans.
, containing the single species C. laeve (formerly C. vulgaris). This was in part due to the stance that Curtis Gates Lloyd
and other mycologists had taken in the early 20th century, believing that the designation of new species was not justifiable due to the existence of intermediate forms in similar habitats and close proximity. In 1970–71, Brodie discovered and reported two variants that differed from C. laeve sufficiently to justify naming them as new species.
-like (obconic) form of the peridia. It differs from Crucibulum laeve in the shape and pink-color of its peridia, as well as its slightly or strongly curved spores (typically 6.5–8 x 11–17 µm). It was found growing on rotten wood and soil in Colombia
by mycologist Gastón Guzmán
.
Peridia are 3–7 mm in diameter x 3–8 mm tall, cup-shaped, short and cylindrical with roughly parallel side walls. The tomentose
exterior surface is tan to yellow when young and whiter in age. Young specimens have an coarsely tomentose epiphragm (membranous cover) that soon disappears. The peridioles are 1–2 mm broad, tan to white in color, disc-shaped, and wrinkled when dry. This species grows on material like twigs, lignin
-rich vegetable debris, wood chips, old matting, or manure.
The immature fruiting body of Crucibulum laeve (technically, the peridium
), is roughly spherical in shape, but in maturity the base is narrowed slightly relative to the top, so that it appears like a cup, or crucible
. The fruiting bodies are usually 5–8 mm tall and almost as wide at the mouth. When young, the mouth is enclosed by a thin membrane called an epiphragm, which is covered with surface hairs. As the fruiting body matures and the fruiting body expands, the epiphragm ruptures, exposing the internal contents. The wall of the fruiting body is made of a single uniform layer of closely interwoven hypha
e (the threadlike filaments that form the mycelium
) roughly 0.25–0.5 mm thick; this wall structure is in contrast to species from the bird's nest fungus genus Cyathus
, which have a distinctly three-layered wall. Young species have a yellowish velvety cover of fine hairs, but this external surface becomes sloughed off and becomes smooth as the fruiting body matures; the color changes to brown, although some old weathered specimens may be bleached grey or dirty white. The inner surface of the fruiting body is smooth and shiny. The cups contain tiny pale ochraceous
or white "eggs," technically termed peridioles, usually 1–2 mm in diameter. In each peridiole is a spore-producing layer of tissue, the hymenium
. This layer is largely composed of basidia (spore-producing cells) mixed with paraphyses
(non-spore producing elements interspersed between basidia). Peridioles are covered by a thin membrane of loosely woven hyphae known as a tunica; separated from the light-colored tunica, the peridioles are black. The peridioles are attached to the inner wall of the peridium by a thin, elastic cord of mycelium, a funiculus, which can be extended at length when moist.
C. laeve has spore
s that are elliptical, hyaline
(translucent), and smooth, with dimensions of 7–10 by 4–6 µm
.
on the outer side and smooth on the inner side, and taper to a narrow base. The peridioles range in width between 0.5–1.25 mm broad. Basidiospores have dimensions of 4–5 by 7–8 µm.
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
in the Nidulariaceae
Nidulariaceae
The Nidulariaceae are a family of fungi in the order Nidulariales. Commonly known as the bird's nest fungi, their fruiting bodies resemble tiny egg-filled birds' nests...
, a family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
of fungi whose fruiting bodies resemble tiny egg-filled bird's nests. Often called "splash cups", the fruiting bodies are adapted for spore
Basidiospore
A basidiospore is a reproductive spore produced by Basidiomycete fungi. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are produced by specialized fungal cells called basidia. In grills under a cap of one common species in the phylum of...
dispersal by using the kinetic energy
Kinetic energy
The kinetic energy of an object is the energy which it possesses due to its motion.It is defined as the work needed to accelerate a body of a given mass from rest to its stated velocity. Having gained this energy during its acceleration, the body maintains this kinetic energy unless its speed changes...
of falling drops of rain. The "eggs" inside the bird's nests (technically known as peridioles) are hard waxy shells containing spores, and tend to stick to whatever nearby herbage they land on, thus increasing the odds of being consumed and dispersed by herbivorous animals. Members of this genus are saprobic, obtaining nutrients from dead organic matter
Organic matter
Organic matter is matter that has come from a once-living organism; is capable of decay, or the product of decay; or is composed of organic compounds...
, and are typically found growing on decayed wood and wood debris. The three known Crucibulum species (C. laeve, C. parvulum, and C. cyathiforme) are distinguished from other genera of the Nidulariaceae by their relatively simple funiculus – a cord of hyphae that connects the peridiole (the "eggs") to the exterior of the bird's nest.
History
Bird's nest fungi were first mentioned by Flemish botanist Carolus Clusius in Rariorum plantarum historia (1601). Over the next couple of centuries, these fungi were the subject of some controversy regarding whether the peridioles were seeds, and the mechanism by which they were dispersed in nature. For example, the French botanist Jean-Jacques Paulet, in his work Traité des champignons (1790–3), erroneously suggested that peridioles were ejected from the fruiting bodies by some sort of spring mechanism.The structure and biology of the genus Crucibulum was better known by the mid-19th century, when the brothers Louis René
Louis René Tulasne
Louis René Tulasne, aka Edmond Tulasne was a French botanist and mycologist who was born in Azay-le-Rideau. He originally studied law at Poitiers, but his interest later turned to botany. As a young man he accompanied botanist Auguste de Saint-Hilaire to South America to study the flora of Brazil...
and Charles Tulasne
Charles Tulasne
Charles Tulasne was a French physician and mycologist who was born in Langeais in the département of Indre-et-Loire. He received his medical doctorate in 1840 and practiced medicine in Paris until 1854. Afterwards he worked with his older brother Louis René Tulasne in the field of mycology...
published a monograph
Monograph
A monograph is a work of writing upon a single subject, usually by a single author.It is often a scholarly essay or learned treatise, and may be released in the manner of a book or journal article. It is by definition a single document that forms a complete text in itself...
on the bird's nest fungi. Subsequently, monographs were written in 1902 by Violet S. White (American species), Curtis Gates Lloyd
Curtis Gates Lloyd
Curtis Gates Lloyd was an American mycologist known for both his research on the Gasteromycetes, as well as his controversial views on naming conventions in taxonomy. He had a herbarium with over 59,000 fungal specimens, and published over a thousand new species of fungi...
in 1906, Gordon Herriot Cunningham
G. H. Cunningham
Gordon Herriot Cunningham C.B.E., F.R.S., was the first New Zealand-based mycologist and plant pathologist. In 1936 he was appointed the first director of the DSIR Plant Diseases Division. Cunningham established the New Zealand Fungal Herbarium, and he published extensively on taxonomy of many...
in 1924 (New Zealand species), and Harold J. Brodie
Harold J Brodie
Harold Johnston Brodie was a Canadian mycologist, known for his contributions to the knowledge of the Nidulariaceae, or bird's nest fungi.-Early life:...
in 1975.
The type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...
for the genus Crucibulum described by the Tulasne brothers was Crucibulum vulgare, an older synonym of the species known today as C. laeve. However, this naming choice was later deemed invalid by rules of fungal nomenclature; the first name validly applied to the species was C. laeve, use by De Candolle, who had based his species upon Nidularia laevis as it appeared in Bulliard's
Jean Baptiste François Pierre Bulliard
Jean Baptiste François Pierre Bulliard was a French physician and botanist....
Histoire des Champignons de la France (Paris, 1791). Kambly and Lee published the first taxonomically
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...
valid description of the genus in 1936. In their 1844 monograph on the Nidulariaceae, the brothers Louis René
Louis René Tulasne
Louis René Tulasne, aka Edmond Tulasne was a French botanist and mycologist who was born in Azay-le-Rideau. He originally studied law at Poitiers, but his interest later turned to botany. As a young man he accompanied botanist Auguste de Saint-Hilaire to South America to study the flora of Brazil...
and Charles Tulasne
Charles Tulasne
Charles Tulasne was a French physician and mycologist who was born in Langeais in the département of Indre-et-Loire. He received his medical doctorate in 1840 and practiced medicine in Paris until 1854. Afterwards he worked with his older brother Louis René Tulasne in the field of mycology...
used the name Crucibulum vulgare, and the species was known by this name until the International Commission on the Taxonomy of Fungi (ICTF) changed the starting-point date for the naming of fungi, and C. vulgare was deemed invalid. The etymology
Etymology
Etymology is the study of the history of words, their origins, and how their form and meaning have changed over time.For languages with a long written history, etymologists make use of texts in these languages and texts about the languages to gather knowledge about how words were used during...
of the specific epithet is derived from the Latin laeve, meaning "smooth".
Description
Crucibulum species have light tanTan (color)
Tan is a pale whiteish, tawny shade of white. The name is derived from tannum used in the tanning of leather.The first recorded use of tan as a shade name in English was in the year 1590....
to cinnamon-colored fruiting bodies
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...
, known as a peridium
Peridium
The peridium is the protective layer that encloses a mass of spores in fungi. This outer covering is a distinctive feature of the Gasteromycetes.-Description:...
, that are cup- or crucible
Crucible
A crucible is a container used for metal, glass, and pigment production as well as a number of modern laboratory processes, which can withstand temperatures high enough to melt or otherwise alter its contents...
-shaped. Depending on the species, the size of the peridium may range from 2–4 tall by 1.5–3 mm wide at the mouth (for C. parvulum) to 5–10 mm tall by 5–8 mm wide (for C. laeve). Viewed microscopically, the wall of the peridium is made of a single layer of tissue, in contrast to the three-layered peridium wall in Cyathus species. The outer surface of the peridium has hypha
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...
e that agglutinate so as to form a texture with visible filaments, a condition known as fibrillose; this outer layers of hairs typically wears off with age to leave a relatively smooth surface.
Mycelium
thumb|right|Fungal myceliaMycelium is the vegetative part of a fungus, consisting of a mass of branching, thread-like hyphae. The mass of hyphae is sometimes called shiro, especially within the fairy ring fungi. Fungal colonies composed of mycelia are found in soil and on or within many other...
, The funicular cord tends to wither away and disappear as the fruiting body ages. Spore
Basidiospore
A basidiospore is a reproductive spore produced by Basidiomycete fungi. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are produced by specialized fungal cells called basidia. In grills under a cap of one common species in the phylum of...
s from Crucibulum species typically have an elliptical or roughly spherical shape, and are thick-walled, transclucent (hyaline
Hyaline
The term hyaline denotes a substance with a glass-like appearance.-Histopathology:In histopathological medical usage, a hyaline substance appears glassy and pink after being stained with haematoxylin and eosin — usually it is an acellular, proteinaceous material...
) or light yellow-brown in color, with dimensions of 5–15 by 5–8 µm
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...
. the spores of C. cyathiforme are notably slightly or strongly curved.
Because the basic fruiting body structure in all genera
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of the Nidulariaceae family is essentially similar, Crucibulum may be readily confused with species of Nidula
Nidula
Nidula is a genus of fungi in the family Nidulariaceae. Like other members of the Nidulariaceae, their fruit bodies resemble tiny egg-filled birds' nests, from which they derive their common name "bird's nest fungi"...
or Cyathus
Cyathus
Cyathus is a genus of fungi in the Nidulariaceae, a family collectively known as the bird's nest fungi. They are given this name since they resemble tiny bird's nests filled with "eggs", structures large enough to have been mistaken in the past for seeds. However, these are now known to be...
, especially older, weathered specimens of Cyathus that may have the hairy ectoperidium worn off. It distinguished from Nidula by the presence of a funiculus, a cord of hyphae attaching the peridiole to the endoperidium. Cyathus differs from genus Crucibulum by having a distinct three-layered wall and a more intricate funiculus.
Peridiole structure
Derived from the GreekGreek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
word peridion, meaning "small leather pouch", the peridiole is the "egg" of the bird's nest. It is a mass of basidiospores and gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...
l tissue enclosed by a hard and waxy outer shell. In Crucibulum, the disc-shaped peridioles are light buff or white; the is due to a persistent layer of tissue surrounding the peridioles, called a tunica. Inside the peridiole is a spore-bearing tissue (the hymenium
Hymenium
The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others some cells develop into sterile cells called cystidia or...
) that is made of spore-bearing cells (basidia), sterile (non-reproductive) structures, and spores.
Peridioles are attached to the fruiting body by a funiculus, a structure of hypha
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...
e that may be differentiated into three regions: the basal piece, which attaches it to the inner wall of the peridium, the middle piece, and an upper sheath, called the purse, connected to the lower surface of the peridiole. In the purse and middle piece is a coiled thread of interwoven hyphae called the funicular cord, attached at one end to the peridiole and at the other end to an entangled mass of hyphae called the hapteron. In Crucibulum species the peridioles is covered by a whitish tunica. The funiculus of Crucibulum species is markedly different from those of Cyathus species: in Crucibulum, the purse is a rounded knob 0.3–0.5 mm wide, attached to the underside of the peridiole. Attaching the purse directly to the wall of the peridium is a stout yellow-grey cord 0.1 mm wide and about 2.5 mm long.
Spore dispersal
John Ray
John Ray was an English naturalist, sometimes referred to as the father of English natural history. Until 1670, he wrote his name as John Wray. From then on, he used 'Ray', after "having ascertained that such had been the practice of his family before him".He published important works on botany,...
in the late 17th century, was tested experimentally by Martin (1927), and more thoroughly by Buller
Arthur Henry Reginald Buller
Arthur Henry Reginald Buller was a British-Canadian mycologist. He is mainly known as a researcher of fungi and wheat rust.- Academic career :...
and Brodie
Harold J Brodie
Harold Johnston Brodie was a Canadian mycologist, known for his contributions to the knowledge of the Nidulariaceae, or bird's nest fungi.-Early life:...
in the 1940s.
Life cycle
The life cycle of Crucibulum, which contains both haploid and diploid stages, is typical of the species of BasidiomycotaBasidiomycota
Basidiomycota is one of two large phyla that, together with the Ascomycota, comprise the subkingdom Dikarya within the Kingdom Fungi...
that can reproduce both asexually (via vegetative
Vegetative reproduction
Vegetative reproduction is a form of asexual reproduction in plants. It is a process by which new individuals arise without production of seeds or spores...
spores), or sexually (with meiosis
Meiosis
Meiosis is a special type of cell division necessary for sexual reproduction. The cells produced by meiosis are gametes or spores. The animals' gametes are called sperm and egg cells....
). Like other wood-decay fungi, this life cycle may be considered as two functionally different phases: the vegetative stage for the spread of mycelia, and the reproductive stage for the establishment of spore-producing structures, the fruiting bodies.
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...
e, pushing out like roots into the rotting wood. These hyphae are homokaryotic
Homokaryotic
Homokaryotic refers to multinucleate cells where all nuclei are genetically identical. It is the antonym of heterokaryotic....
, containing a single nucleus in each compartment; they increase in length by adding cell-wall material to a growing tip. As these tips expand and spread to produce new growing points, a network called the mycelium develops. Mycelial growth occurs by mitosis
Mitosis
Mitosis is the process by which a eukaryotic cell separates the chromosomes in its cell nucleus into two identical sets, in two separate nuclei. It is generally followed immediately by cytokinesis, which divides the nuclei, cytoplasm, organelles and cell membrane into two cells containing roughly...
and the synthesis of hyphal biomass. When two homokaryotic hyphae of different mating compatibility groups
Mating type
Mating types occur in eukaryotes that undergo sexual reproduction via isogamy. Since the gametes of different mating types look alike, they are often referred to by numbers, letters, or simply "+" and "-" instead of "male" and "female." Mating can only take place between different mating...
fuse with one another, they form a dikaryotic
Dikaryon
Dikaryon is from Greek, di meaning 2 and karyon meaning nut, referring to the cell nucleus.The dikaryon is a nuclear feature which is unique to some fungi, in which after plasmogamy the two compatible nuclei of two cells pair off and cohabit without karyogamy within the cells of the hyphae,...
mycelia in a process called plasmogamy
Plasmogamy
Plasmogamy is a stage in the sexual reproduction of fungi. In this stage, the cytoplasm of two parent mycelia fuse together without the fusion of nuclei, as occurs in higher terrestrial fungi. After plasmogamy occurs, the secondary mycelium forms. The secondary mycelium consists of dikaryotic...
. Prerequisites for mycelial survival and colonization a substrate (like rotting wood) include suitable humidity and nutrient availability.Crucibulum laeve is saprobic, so mycelial growth in rotting wood is made possible by the secretion of enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
s that break down complex polysaccharide
Polysaccharide
Polysaccharides are long carbohydrate molecules, of repeated monomer units joined together by glycosidic bonds. They range in structure from linear to highly branched. Polysaccharides are often quite heterogeneous, containing slight modifications of the repeating unit. Depending on the structure,...
s (such as cellulose
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....
and lignin
Lignin
Lignin or lignen is a complex chemical compound most commonly derived from wood, and an integral part of the secondary cell walls of plants and some algae. The term was introduced in 1819 by de Candolle and is derived from the Latin word lignum, meaning wood...
) into simple sugars that can be used as nutrients.
After a period of time and under the appropriate environmental conditions, the dikaryotic mycelia may enter the reproductive stage of the life cycle. Fruiting body formation is influenced by external factors such as season (which affects temperature and air humidity), nutrients and light. As fruiting bodies develop they produce peridioles containing the basidia upon which new basidiospores are made. Young basidia contain a pair of haploid sexually compatible nuclei which fuse, and the resulting diploid fusion nucleus undergoes meiosis to produce basidiospore
Basidiospore
A basidiospore is a reproductive spore produced by Basidiomycete fungi. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are produced by specialized fungal cells called basidia. In grills under a cap of one common species in the phylum of...
s, each containing a single haploid nucleus. The dikaryotic mycelia from which the fruiting bodies are produced is long lasting, and will continue to produce successive generations of fruiting bodies as long as the environmental conditions are favorable.
Development
The initial studies on the development of the fruiting bodies in Crucibulum were performed by the brothers Tulasne (1844), Sachs (1855), DeBary (1866), Eidam (1877), and Walker (1920). Collectively, these early researchers determined that basidiosporeBasidiospore
A basidiospore is a reproductive spore produced by Basidiomycete fungi. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are produced by specialized fungal cells called basidia. In grills under a cap of one common species in the phylum of...
s are produced on club-shaped basidia which line the internal cavity of the peridiole. Basidia typically have 4 spores, attached by a short projection (a sterigma
Sterigma
thumb|right|The sterigmata is the slender extension that connects the spore to the basidia .A sterigma is an extension of the basidium consisting of a basal filamentous part and a slender projection with a spore at the tip...
); after being detached from the basidia the spores migrate towards the center of the peridiole concurrently with the collapse and gelatinization of the underlying tissues.
Bioactive compounds
Chemical structure
A chemical structure includes molecular geometry, electronic structure and crystal structure of molecules. Molecular geometry refers to the spatial arrangement of atoms in a molecule and the chemical bonds that hold the atoms together. Molecular geometry can range from the very simple, such as...
related to benzofuran
Benzofuran
Benzofuran is the heterocyclic compound consisting of fused benzene and furan rings. This colourless liquid is a component of coal tar. Benzofuran is the "parent" of many related compounds with more complex structures. For example, psoralen is a benzofuran derivative that occurs in several...
and chromene, molecules that contain cyclic amide
Amide
In chemistry, an amide is an organic compound that contains the functional group consisting of a carbonyl group linked to a nitrogen atom . The term refers both to a class of compounds and a functional group within those compounds. The term amide also refers to deprotonated form of ammonia or an...
or lactone
Lactone
In chemistry, a lactone is a cyclic ester which can be seen as the condensation product of an alcohol group -OH and a carboxylic acid group -COOH in the same molecule...
five-membered ring structures; these compounds are unique to this species. Salfredin B11 was first identified in 1995, while later research confirmed the presence of additional salfredin-type metabolites
Secondary metabolite
Secondary metabolites are organic compounds that are not directly involved in the normal growth, development, or reproduction of an organism. Unlike primary metabolites, absence of secondary metabolities does not result in immediate death, but rather in long-term impairment of the organism's...
. These compounds are inhibitors
Enzyme inhibitor
An enzyme inhibitor is a molecule that binds to enzymes and decreases their activity. Since blocking an enzyme's activity can kill a pathogen or correct a metabolic imbalance, many drugs are enzyme inhibitors. They are also used as herbicides and pesticides...
of aldose reductase
Aldose reductase
Aldose reductase is an NADPH-dependent oxidoreductase that catalyzes the reduction of a variety of aldehydes and carbonyls, including monosaccharides...
, an enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
that has been implicated in the formation of cataract
Cataract
A cataract is a clouding that develops in the crystalline lens of the eye or in its envelope, varying in degree from slight to complete opacity and obstructing the passage of light...
s in advanced stages of diabetes mellitus
Diabetes mellitus
Diabetes mellitus, often simply referred to as diabetes, is a group of metabolic diseases in which a person has high blood sugar, either because the body does not produce enough insulin, or because cells do not respond to the insulin that is produced...
. The salfredin compounds may have therapeutic use in the treatment of this disorder.
Habitat and distribution
Like other bird's nest fungi, Crucibulum species are saprobic and derive their nutrients from decomposing organic matterOrganic matter
Organic matter is matter that has come from a once-living organism; is capable of decay, or the product of decay; or is composed of organic compounds...
. They are typically found growing on wood and woody debris such as stems, twigs, wood chips, old nut shells, and old matting; they are sometimes found on "dried manure cakes". Brodie notes (of C. laeve) they are "never" found on soil or large logs. C. parvulum has been found on the roots and stems of old or dead dry land plants such as Juniperus horizontalis
Juniperus horizontalis
Juniperus horizontalis is a low-growing shrubby juniper native to northern North America, throughout most of Canada from Yukon east to Newfoundland, and in the United States in Alaska, and locally from Montana east to Maine, reaching its furthest south in Wyoming and northern Illinois.It lives up...
and Artemisa
Artemisa
Artemisa is a municipality and city in Cuba, formerly part of La Habana Province.According to a law approved by the Cuban National Assembly in August 2010, Artemisa became the capital city of the newly formed Artemisa Province, which comprises 8 municipalities of the former La Habana Province and 3...
species.
C. laeve, the most well-known species of Crucibulum, is a temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...
-zone species with a circumpolar distribution. It has been collected in most European countries and the Canary Islands
Canary Islands
The Canary Islands , also known as the Canaries , is a Spanish archipelago located just off the northwest coast of mainland Africa, 100 km west of the border between Morocco and the Western Sahara. The Canaries are a Spanish autonomous community and an outermost region of the European Union...
; in North America it has been found from Alaska to Mexico, while South American locations include Chile and Tierra del Fuego
Tierra del Fuego
Tierra del Fuego is an archipelago off the southernmost tip of the South American mainland, across the Strait of Magellan. The archipelago consists of a main island Isla Grande de Tierra del Fuego divided between Chile and Argentina with an area of , and a group of smaller islands including Cape...
. It has also been found in Australia, Iceland, Japan, and New Zealand. C. parvulum has a primarily North American distribution, having been found in Alaska
Alaska
Alaska is the largest state in the United States by area. It is situated in the northwest extremity of the North American continent, with Canada to the east, the Arctic Ocean to the north, and the Pacific Ocean to the west and south, with Russia further west across the Bering Strait...
, southern Alberta
Alberta
Alberta is a province of Canada. It had an estimated population of 3.7 million in 2010 making it the most populous of Canada's three prairie provinces...
Badlands
Badlands
A badlands is a type of dry terrain where softer sedimentary rocks and clay-rich soils have been extensively eroded by wind and water. It can resemble malpaís, a terrain of volcanic rock. Canyons, ravines, gullies, hoodoos and other such geological forms are common in badlands. They are often...
, the Canadian Rocky Mountains
Rocky Mountains
The Rocky Mountains are a major mountain range in western North America. The Rocky Mountains stretch more than from the northernmost part of British Columbia, in western Canada, to New Mexico, in the southwestern United States...
and in semi-deserts of Idaho
Idaho
Idaho is a state in the Rocky Mountain area of the United States. The state's largest city and capital is Boise. Residents are called "Idahoans". Idaho was admitted to the Union on July 3, 1890, as the 43rd state....
; in 2004 it was collected in China. C. cyathiforme is only known from Colombia—where it was discovered growing at an elevation of nearly 7000 feet (2146 metres)— and Armenia
Armenia
Armenia , officially the Republic of Armenia , is a landlocked mountainous country in the Caucasus region of Eurasia...
.
Edibility
Species in the Nidulariaceae family, including Crucibulum, are considered inedible, as they are "not sufficiently large, fleshy, or odorous to be of interest to humans as food". However, there have not been reports of poisonous alkaloidAlkaloid
Alkaloids are a group of naturally occurring chemical compounds that contain mostly basic nitrogen atoms. This group also includes some related compounds with neutral and even weakly acidic properties. Also some synthetic compounds of similar structure are attributed to alkaloids...
s or other substances considered toxic to humans.
Species
Until the 1970s Crucibulum was thought to be monotypicMonotypic
In biology, a monotypic taxon is a taxonomic group with only one biological type. The term's usage differs slightly between botany and zoology. The term monotypic has a separate use in conservation biology, monotypic habitat, regarding species habitat conversion eliminating biodiversity and...
, containing the single species C. laeve (formerly C. vulgaris). This was in part due to the stance that Curtis Gates Lloyd
Curtis Gates Lloyd
Curtis Gates Lloyd was an American mycologist known for both his research on the Gasteromycetes, as well as his controversial views on naming conventions in taxonomy. He had a herbarium with over 59,000 fungal specimens, and published over a thousand new species of fungi...
and other mycologists had taken in the early 20th century, believing that the designation of new species was not justifiable due to the existence of intermediate forms in similar habitats and close proximity. In 1970–71, Brodie discovered and reported two variants that differed from C. laeve sufficiently to justify naming them as new species.
C. cyathiforme
The specific epithet of this species refers to the vase-like or CyathusCyathus
Cyathus is a genus of fungi in the Nidulariaceae, a family collectively known as the bird's nest fungi. They are given this name since they resemble tiny bird's nests filled with "eggs", structures large enough to have been mistaken in the past for seeds. However, these are now known to be...
-like (obconic) form of the peridia. It differs from Crucibulum laeve in the shape and pink-color of its peridia, as well as its slightly or strongly curved spores (typically 6.5–8 x 11–17 µm). It was found growing on rotten wood and soil in Colombia
Colombia
Colombia, officially the Republic of Colombia , is a unitary constitutional republic comprising thirty-two departments. The country is located in northwestern South America, bordered to the east by Venezuela and Brazil; to the south by Ecuador and Peru; to the north by the Caribbean Sea; to the...
by mycologist Gastón Guzmán
Gastón Guzmán
Gastón Guzmán is a Mexican mycologist and anthropologist. He is considered the world's foremost authority on the genus Psilocybe.He was born in Xalapa, Veracruz, in 1932....
.
C. laeve
Tomentose
Tomentose is a term used to describe plant hairs that are flattened and matted, forming a woolly coating known as tomentum. Often the hairs are silver or gray-colored...
exterior surface is tan to yellow when young and whiter in age. Young specimens have an coarsely tomentose epiphragm (membranous cover) that soon disappears. The peridioles are 1–2 mm broad, tan to white in color, disc-shaped, and wrinkled when dry. This species grows on material like twigs, lignin
Lignin
Lignin or lignen is a complex chemical compound most commonly derived from wood, and an integral part of the secondary cell walls of plants and some algae. The term was introduced in 1819 by de Candolle and is derived from the Latin word lignum, meaning wood...
-rich vegetable debris, wood chips, old matting, or manure.
The immature fruiting body of Crucibulum laeve (technically, the peridium
Peridium
The peridium is the protective layer that encloses a mass of spores in fungi. This outer covering is a distinctive feature of the Gasteromycetes.-Description:...
), is roughly spherical in shape, but in maturity the base is narrowed slightly relative to the top, so that it appears like a cup, or crucible
Crucible
A crucible is a container used for metal, glass, and pigment production as well as a number of modern laboratory processes, which can withstand temperatures high enough to melt or otherwise alter its contents...
. The fruiting bodies are usually 5–8 mm tall and almost as wide at the mouth. When young, the mouth is enclosed by a thin membrane called an epiphragm, which is covered with surface hairs. As the fruiting body matures and the fruiting body expands, the epiphragm ruptures, exposing the internal contents. The wall of the fruiting body is made of a single uniform layer of closely interwoven hypha
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...
e (the threadlike filaments that form the mycelium
Mycelium
thumb|right|Fungal myceliaMycelium is the vegetative part of a fungus, consisting of a mass of branching, thread-like hyphae. The mass of hyphae is sometimes called shiro, especially within the fairy ring fungi. Fungal colonies composed of mycelia are found in soil and on or within many other...
) roughly 0.25–0.5 mm thick; this wall structure is in contrast to species from the bird's nest fungus genus Cyathus
Cyathus
Cyathus is a genus of fungi in the Nidulariaceae, a family collectively known as the bird's nest fungi. They are given this name since they resemble tiny bird's nests filled with "eggs", structures large enough to have been mistaken in the past for seeds. However, these are now known to be...
, which have a distinctly three-layered wall. Young species have a yellowish velvety cover of fine hairs, but this external surface becomes sloughed off and becomes smooth as the fruiting body matures; the color changes to brown, although some old weathered specimens may be bleached grey or dirty white. The inner surface of the fruiting body is smooth and shiny. The cups contain tiny pale ochraceous
Ochre
Ochre is the term for both a golden-yellow or light yellow brown color and for a form of earth pigment which produces the color. The pigment can also be used to create a reddish tint known as "red ochre". The more rarely used terms "purple ochre" and "brown ochre" also exist for variant hues...
or white "eggs," technically termed peridioles, usually 1–2 mm in diameter. In each peridiole is a spore-producing layer of tissue, the hymenium
Hymenium
The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others some cells develop into sterile cells called cystidia or...
. This layer is largely composed of basidia (spore-producing cells) mixed with paraphyses
Paraphyses
Paraphyses are part of the fertile spore-bearing layer in certain fungi. More specifically, paraphyses are sterile filamentous hyphal end cells composing part of the hymenium of Ascomycota and Basidiomycota interspersed among either the asci or basidia respectively, and not sufficiently...
(non-spore producing elements interspersed between basidia). Peridioles are covered by a thin membrane of loosely woven hyphae known as a tunica; separated from the light-colored tunica, the peridioles are black. The peridioles are attached to the inner wall of the peridium by a thin, elastic cord of mycelium, a funiculus, which can be extended at length when moist.
C. laeve has spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...
s that are elliptical, hyaline
Hyaline
The term hyaline denotes a substance with a glass-like appearance.-Histopathology:In histopathological medical usage, a hyaline substance appears glassy and pink after being stained with haematoxylin and eosin — usually it is an acellular, proteinaceous material...
(translucent), and smooth, with dimensions of 7–10 by 4–6 µm
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...
.
C. parvulum
This species is characterized by its very small peridia (dimensions 1.5–3 mm wide at the mouth x 2–4 mm tall) with a color that may range from white to grey to pale buff, but never yellow – helping distinguish it from C. laeve. The peridia are obconic, thin-walled (150–180 µm at the lip, approximately 300 µm thick at the edge of the lip), tomentoseTomentose
Tomentose is a term used to describe plant hairs that are flattened and matted, forming a woolly coating known as tomentum. Often the hairs are silver or gray-colored...
on the outer side and smooth on the inner side, and taper to a narrow base. The peridioles range in width between 0.5–1.25 mm broad. Basidiospores have dimensions of 4–5 by 7–8 µm.