View allAll Photos Tagged differences,
In 2010, the main difference was the presence of Ardrossan Indoor Bowling Club, built at the left, built shortly after the photo was taken. The Town Hall was built in 1851 by Duncan Graham from the north of Scotland and was known as Graham’s Castle. From 1893, it was owned by Archibald Russell and his heirs. In 1920, it was bought by the Ardrossan Dry Dock and Shipbuilding Company Limited who refurbished it as Castlecraigs Recreation Club which opened on 26 February 1921. In 1927, the Masons bought the building and let out the ancillary rooms and tennis courts. During the second world war between 1939 and 1945, the Royal Navy requisitioned Castlecraigs as a barracks. It was subsequently managed by Ardrossan Town Council and known as the Town Hall. In the 1970s, North Ayrshire Council took over the building, added an extension to the right and called it Ardrossan Civic Centre.
Photograph contributed by Tom McGrattan
Website: www.ardrossanphotographs.net/
For more information on the Yesterd@ys project, please visit Our Website, or email us at NAHeritage@North-Ayrshire.gov.uk
DISCLAIMER
All archival images on this website have been made available by The North Ayrshire Council in good faith for reference and/or educational purposes only and without intent to breach any proprietary rights which may subsist in the work. Images may not be printed, copied, distributed, published or used for any commercial purposes without the prior written consent of the individual or body which holds such rights. Should any alleged breach of proprietary rights be brought to the attention of The North Ayrshire Council, relevant material will be removed from the website with immediate effect.
The North Ayrshire Council is not responsible for the content, reliability or availability of external websites and cannot be held liable for any loss or damage to the user, of whatever kind, arising either directly or indirectly from use of same. Listing should not be taken as an endorsement of any kind and in particular, of views expressed within any such site.
113in2013 16 - Spot the Difference.
There are nine differences between the top photo and the bottom photo. Can you find them all? (Artist Ann Swanson Gross)
I am building a small visualization tool to look at the similarities and differences between two articles published in October about head injuries and the NFL:
"Game Brain" by Jeanne Marie Laskas - Oct. 10, 2009
www.gq.com/sports/profiles/200909/nfl-players-brain-demen...
"Offensive Play" by Malcolm Gladwell - Oct. 19, 2009
www.newyorker.com/reporting/2009/10/19/091019fa_fact_glad...
These are some early outputs from the system.
The tree had a really catching difference! Some souls are exactly in the same way! One will feel so jealous to see such people...
This image of was taken in the Mount Field National Park in Tasmania, Australia. There are three falls in the park including Russell Falls, Lady Baron Falls and Horseshoe Falls. I now have images of these 3 falls and other parts of the park which I will be gradually posting in this photostream.
I rate this National Park as the most photogenic place I have been to in Tasmania. Next time I will be focussing on the Talls Trees Circuit and macro images of this wonderful rainforest environment.
Geology
The park lies to the west of the Derwent Graben, formed during the mid-Tertiary. Outcropping to the west of the park, and underlying it as basement rock, are strongly folded older successions of Ordovician and Siluro-Devonian sediments including the Gordon limestone and its equivalents. Jurassic dolerite is ubiquitous above about 760 metres, with Triassic and Permian sediments outcropping at lower altitudes.
The Triassic and Jurassic rocks of the park and reserves show strong Gondwanic links with those of the Transantarctic Mountains in Antarctica. The Triassic sequence of sedimentary rocks is very uniform, non-marine in origin and contains evidence of lacustrine and fluvial conditions with basic intrusive material.
The basalt dyke visible near Mt Bridges above Lake Seal is evidence of a recent intrusion in the faulted dolerite. The Jurassic dolerite (170 million years old) provides a firm link with Antarctica being of identical age.
Lady Barron Falls, Horseshoe Falls and Russell Falls are composed of horizontally bedded marine Permian siltstone benches. The vertical faces of the waterfalls are composed of the more resistant sandstone layers along vertical joint.
Aboriginal Heritage
Limited archaeological surveys in the park have shown that Aborigines used the land and waters of the park; more extensive surveys of the nearby Florentine Valley have shown Aboriginal occupation of over 30,000 years.
To date there have been no systematic archaeological surveys conducted in the park and reserves. Two Aboriginal sites have been identified inside the park boundaries, located near Lake Fenton and Lake Dobson.
The park fell within the territory of the Big River people, who occupied territory ranging from the Great Western Tiers to Mount Wellington. The Pangerninghe band located at the junction of the Derwent and Clyde Rivers near the present-day township of Hamilton, were the closest band to the park and reserves area.
European History
Early reports by white settlers noted the presence of Aboriginal people in the Russell Falls area. One such report dated 1827 noted an Aboriginal party of some 60 persons engaged in a skirmish with landowner Charles Abbott's servants, until the overseer arrived and "prudently settled all differences by the distribution of some provision amongst them" (The Tasmanian, 16 November 1827 p.2 col.1.).
From the 1830s, trappers and snarers worked the high country around Mt Field, coming in from Montos Marsh (now Ellendale). Bushrangers and escaped convicts were known to have hidden out in the country around Bushy Park in the 1840s, trapping and taking advantage of local farms such as Fenton Forest.
Prior to 1910, the only access to the high country was by pack track from Ellendale. In 1869, the eminent botanist Baron von Mueller visited Mt Field East on a week-long collecting trek, guided by the Rayner brothers, local trappers. He described the snow gum (Eucalyptus coccifera), the urn gum (E. urnigera), the cider gum (E. gunnii) and cushion plants (Donatia novaezelandie) from the meadows around Lake Fenton. The botanist Leonard Rodway also explored the area from the 1850s.
The first non-Aboriginal to encounter Russell Falls was a settler named Browning in 1856. The falls became known as Brownings Falls until about 1884 when confusion caused them to be referred to as Russell Falls, and by the turn of the century, the name was firmly established.
The original Russell Falls, named for a member of an exploration party in the Derwent Valley, was actually located on the Tyenna River, which was previously known as Russell Falls River. Frodsham surveyed the area in 1884, and the Falls Reserve was proclaimed in 1885.
Source: www.parks.tas.gov.au/index.aspx?base=3601
FOOTNOTE.
Apart from cropping and brightness/contrast adjustments, all of my images are presented as they were when the shutter closed, unless otherwise stated
I'm now subtracting 5 in the 5th position from the left. This is done in 3 steps: subtract 50, add 40, add 5. In this step I add 40 by adding 4 in the 4th position from the left.
© Milan Cvetanovic
All rights reserved!
I rarely do posed shots at events, I make exception when the subjects ask for a photo themselves, ask to be photographed. This is such an occasion, captured at the opening day of the 2025 Ballinasloe Horse Fair (County Galway, Ireland), the oldest horse fair in Europe, held since the first half of the 1700s. All horse owners and buyers, horse riders and lovers of horses from the region and the whole island gather during this week in Ballinasloe. It is a traditional meeting place of the Irish Travellers as well, who resemble in their lifestyle to the Roma (Gypsies), yet there are not genetically connected with them. The Travellers diverged from the Irish about 500 years ago. www.nature.com/articles/srep42187
Much thanks for your kind support!
20170516GraduationCapGownFrederickRoyster
Major: Interdisciplinary Studies, Concentration in Education, Minor in Special Education
Henderson Nc
What is your best memory of WSSU?
My best memory of WSSU would be Homecoming! All the support, love and positive energy from alumni, students, faculty and staff is amazing! Seeing thousands of people coming together, bring their kids and relatives to brag about how awesome WSSU is priceless. To celebrate this wonderful institution through donations, attendance at athletic events as well as fellowshipping on the yard is phenomenal!
What do you love about WSSU?
The one thing I love about WSSU is the family oriented vibe I receive from not only the faculty and staff, but also from the students? There is not a moment that goes by each and every day where I do not see the genuine vibe and care our students have for one another. Wither it is student leaders encouraging other students to get more involved with student activities or students coming together to walk to the football or basketball games to support athletics or senate meetings, together we act as one.
What do you plan on doing after you graduate?
My plans after graduation are to further my education to receive my masters in the field of education. With this goal I plan to teach at a Title 1 school in the Forsyth County region to help leave a legacy in the community of Winston Salem and embrace my love and the importance of my HBCU.
How has WSSU prepared you for the future?
Being that WSSU is a Liberal Arts Institution is has prepared me with the knowledge and skill set in different content areas where I will be able to use that background knowledge and the fundamental skill set to adapt to any job I come across in the future in life.
How has WSSU made a difference in your life?
WSSU has made a huge difference in my life. Getting involved in different organizations such as Greek Life, Housing and Resident Life and Student Government Association has taught me to believe in myself. Getting involved in campus life has developed my communication and social skills. It has taught me how important your personal brand is and taught me the importance of networking and getting out of my comfort zone. If I did not have individuals at this this institution to push and help motivate me, I would have never revealed my full potential. And for that I am proud to be an alum of the Winston Salem State University!
The two editions of the Megabot pose for a special photo. You can see a slight color difference, though the camera flash dulled the changes. To the left is the original, and on the right is the reissue. The 2nd edition is missing paint detail on the lower front hatch and also missing the Void symbol on the chest panel. The most noticeable color difference lies in the hand pieces, as the reissue has flat gray while the original has a blue/gray mixture. Be sure to view the full size photo for the best detail!
Which is your favorite?
Canon 6D Mark II with Sigma 24-70mm lens.
Captured during the "On Fire" weekend of May 25-26. Sunday June 9th there are elections for the Parliaments.
Together with Sarah we took several pics of this event @ Leuven.
After the speeches the president of the PVDA took a group selfie with other candidates and the crowd.
Not my best technical photo (because of the difference in light on stage and beyond), but it surely represents the atmosphere of the day.
If you know what you're looking at you'll spot the difference straight away!
Tina, my trusty coffee machine in work, has developed a bit of a continence problem. With a full tank she's taken to pissing out a bit of water overnight.
The culprit is this rubber seal. It sits underneath the water tank and forms the seal that prevents leakage. The one on the right is the old one. Its inner wall is flat, like a polo mint. The one on the left is the replacement - its walls are tapered so the hole at the bottom is smaller than the hole at the top.
I wasn't sure about getting hold of parts, but it turns out that the Spidem Trevi Chiara is a Saeco Vienna in drag so parts are easy to come by, and the beandoctor sorted me out a replacement for a fiver with postage.
Happy days.
Differences, Beauty, Sadness, Love, Life, Death, Dreams, Desires,
Hopes, Better days, Different lives, hands, Hug, Tears,
Light, Neighberhood, Bonding history, Appertunities, Failurs,
Agony, Passion, Heart, Wisdom, You, Myself,
Us, We, Them, Future!
this is what I see In Another's Eyes - from my eyes while looking at you.
some of it is what I Wish not to see.
some of it is A wish with A prayers to be heard.
Now,
while you're looking at me, Do you feel the same?
I've this set included an unique minifigure Nexo knight.
Here you can see some differents details on the legs and on the torso from the other knight in the sets 70310, 70311, 30373 and 30371.
Original picture by me
Small things can make a whole big difference.
I was sorry to hear today that a comment written by a friend on my Flickr portfolio disappeared before, I even saw it.
I was also very sorry to discover today, that at least 2 friendship links at my FB disappeared too.
I did not delete any of those, and it can be that more were deleted unfortunately and not by myself. I believe this was done deliberately.
I would say sorry in advance.. And surely I can't search for any kind of "missing" on daily base. I only sometimes check my friends links.
All I could say, I never deleted a friend.
I vote for a better world... and better friendships.
Thanks for visiting.
Copyrights (c) Nira Dabush.
Devotees of the www.flickr.com/photos/nlireland/ photosream might think that this picture by Waterford photographer A.H. Poole looks familiar. But this one, from the UK National Archives at Kew, differs in several ways (I spent a considerable time yesterday wearing white cotton gloves squinting through a magnifying glass at the 1910 print).
Simliarities;
- Poole was in the same position for both photos (the lamppost aligns with the T of Waterford on both photos),
- and both photos were subject to the same 'photoshopping' (the ends of the shafts removed and the W of Waterford restored)
Differences
- The Kew picture has the van in the centre, rather than to the left
- The people (other than the tea ladies) and cattle are different (or not obviously the same as far as I could judge)
- the NLI photo has some liquid manure (I presume) trickling towards the van, suggesting it was taken after the Kew one.
- The board advertising Lipton's tea looks as if it is leaning against the steps. and barely visible.
- the printer of the Kew photo has de-emphasised the buildings in the background.
What a difference a few hours makes in autumn! Most of those fallen red acer leaves in our woodland lower garden fell overnight!
After a wet summer in the West Midlands many of the autumn foliage colours have been intense this year. However recent air frosts, wind and rain have brought an abrupt end to some of their displays!
Charity Garden Open days in 2013 include Winter (February 24), Late spring, Summer and Autumn.
Please do not use this image without my prior consent.
Morgenthal Frederics PURO Eyeglasses. Lens 39mm (with lens holder) D.B.L. 22mm
PURO is a unique, simple design eyeglasses. Titanium frame with acetate lens holder. The Ti frame comes in pewter, polished chrome, gummetal, bronze finish. Lens holder comes in White, Black, Brown, Tortoise and Horn. PURO are sold with or without lens holders with price difference. The acetate piece is not a decoration piece that mount on the Ti frame. The lens actually set in it instead of Ti frame itself. It is very cool looking and no need to worry that they will fell out. PURO is not perfectly round or oval.
As pretty as it is, I had THE WORSE experience EVER with PURO. It was purchased at Morgenthal Frederics at 59th Street in Time Warner Center building. Sale was helpful cuz I had intend of buying. At the time, I wasn't sure to go for acetate or not and was told the lens holder can be purchase separately if I change my mind. So I went for the Ti frame only. The frame was around $420 and another $400 for the lens. The $400 lens doesn't include all the coating and 4 times more than what I usually pay for. After owning it for 3 month, I found out that the frame is not perfect round or oval. The imperfect roundness bother my mind greatly. I thought the acetate lens holder would help to notice the frame shape so I thought I'll pay them a visit.
I went back to the store I bought it from and told the sales I would like to purchase the lens holder and would nice if they can insert it for me. It was different sales this time and he told me "NO, we don't sell them separately." I told them I was inform so and how can I get a lens holder? "Sir, you have to purchase a brand new frame with it." I told them the idea was ridiculous, cuz if the told me so the first place, I would made different choice. "Sir, we can't help you unless you buy a brand new frame." I left the store ANGRY and thought she don't know what she is talking about, let me went a different store and see what they say. Some one gotta make some sense right?!
I went to their store on the 63rd & Madison with the same request. Sales ask me where my frame originally from and call the Time Warner store to confirm. While he was on the phone, he was judging me through his eyes and told me "NO. They said you were just at Time Warner Store and they have told you everything you need to know. I am sorry". I was FURIOUS this time.
Still not giving it up, thanks god I got unlimited subway pass. I went to 74th Street & Madison store. I told them I went through 2 stores already if they wanna call and confirm, be my guest. I told them the whole story and what 2 stores were tell me and how I am furious before I enter this store and hoping they can give me a better answer. They respond. "We can give you the lens holder for free, but you have to buy a set of brand lens from us." (which was $400) I asked "why, optician can just re-shape it". They respond, "The lens is too thin, they will break." I told them, "why don't you sell me the lens holder, and I will ask my optician." They say, "sorry, we can't do that. Not unless you buy a set of brand new lens. What an unbelievable bunch of assholes!
At last I went to their last store in Manhattan located at W. Broadway in SoHo. I told spoke with the manager, told them the 3 stores responses. I let him decided if anything make any sense. "They might misunderstood with your request. You know what, I'll give the lens holder to you for free. We want our customer to be happy." Took me to go through all stores with anger. All, because their sales misinform their customer.
I took the lens holder to Abe, my optician in Chinatown. I ask him if he can reshape my current lens into the lens holder. Took Abe 5 min. and lens set in the holder. "the lens is made of plastic not glass, why they think it will break?" I took the frame back, scratch & Spot less, PERFECT. I ask Abe, "Can you check the lens quality and coating, how much you will charge to match it." Abe check it and responded, "$120, for you, I'll charge $100. This lens doesn't have all the coating you usually have on. $100 you got everything." I ask him if this lens is higher end, thinner, better in any other way? "No, it is the same thing we are using here. It is not better or special." I did not respond. I did not tell Abe about Morgenthal Frederics charge $400 dollars for them with less coating. They absolutely made me feel like a FOOL.
Everything above is 100% true. Morgenthal Frederics products no doubt are well made but people working for the company seem too money hungry. I told everyone what you guys have done so they can have 2nd thoughts before buying Morgenthal Frederics products. Morgenthal Frederics sure gave me the worse shopping experience. Now, whenever people search for Morgenthal Frederics online, my review will show up with it. I will NEVER buy another pair from you guys and I hope this review will HURT your company deeply.
Customers are the reasons why your company is still standing. Remember it well and treat them good. WORDS do spread. Reputation has to earn it by SERVICE.
dannyman.toldme.com/2010/06/28/charles-babbages-differenc...
I tingled inside at the sight of this beauty. Charles Babbage was a genius who designed a mechanical, base-10 computing device way before the modern computer era. His vision was never built: it was just too hard and expensive and plain old ahead of its time. Finally, in the 1980s, this computer was built based on Babbage's old designs. A beautiful brass hand-cranked calculating machine! For a modern computer geek this is not unlike seeing a dinosaur brought to life.
Produced this for my self sanity and anybody else interested. The blanks in the 'Pro account - New' are the same as those stated in the 'Free Account'.
The 'Pro account - New' was populated by using the details from the upgrade button (not sure how it is an upgrade when I am already a pro account holder, or was).
I have to say at this point I would be making a contractual arrangement By clicking “UPGRADE NOW” you agree to receive Flickr Pro from Yahoo! Inc. under the Yahoo! Inc. The agreement on this page does not specify that the Pro Account will include the products and services as listed in 'Free Account'.
I believe the actions of Yahoo to be unethical in consideration of those with Pro Account contracts, the question is should Yahoo - Flickr continue to deliver the same services and products you agreed to when you previously made you contract or do you want the addition storage benefits.
As for the home page and interface changes compared to the other services, I still think Flickr has richer content on word searching and the community interactions (if there is anyone left in Flickr land).
If you had a free account previously and you're jumping around because of the home page change, interesting to see how it will cost to jump ship to another provider, what I seen so far to get something decent £6.95 per month sounds a lot more than zero.
A fungus (pl.: fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as one of the traditional eukaryotic kingdoms, along with Animalia, Plantae and either Protista or Protozoa and Chromista.
A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is chitin in their cell walls. Fungi, like animals, are heterotrophs; they acquire their food by absorbing dissolved molecules, typically by secreting digestive enzymes into their environment. Fungi do not photosynthesize. Growth is their means of mobility, except for spores (a few of which are flagellated), which may travel through the air or water. Fungi are the principal decomposers in ecological systems. These and other differences place fungi in a single group of related organisms, named the Eumycota (true fungi or Eumycetes), that share a common ancestor (i.e. they form a monophyletic group), an interpretation that is also strongly supported by molecular phylogenetics. This fungal group is distinct from the structurally similar myxomycetes (slime molds) and oomycetes (water molds). The discipline of biology devoted to the study of fungi is known as mycology (from the Greek μύκης mykes, mushroom). In the past mycology was regarded as a branch of botany, although it is now known that fungi are genetically more closely related to animals than to plants.
Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil or on dead matter. Fungi include symbionts of plants, animals, or other fungi and also parasites. They may become noticeable when fruiting, either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient cycling and exchange in the environment. They have long been used as a direct source of human food, in the form of mushrooms and truffles; as a leavening agent for bread; and in the fermentation of various food products, such as wine, beer, and soy sauce. Since the 1940s, fungi have been used for the production of antibiotics, and, more recently, various enzymes produced by fungi are used industrially and in detergents. Fungi are also used as biological pesticides to control weeds, plant diseases, and insect pests. Many species produce bioactive compounds called mycotoxins, such as alkaloids and polyketides, that are toxic to animals, including humans. The fruiting structures of a few species contain psychotropic compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant pathogens of humans and other animals. Losses of crops due to fungal diseases (e.g., rice blast disease) or food spoilage can have a large impact on human food supplies and local economies.
The fungus kingdom encompasses an enormous diversity of taxa with varied ecologies, life cycle strategies, and morphologies ranging from unicellular aquatic chytrids to large mushrooms. However, little is known of the true biodiversity of the fungus kingdom, which has been estimated at 2.2 million to 3.8 million species. Of these, only about 148,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans. Ever since the pioneering 18th and 19th century taxonomical works of Carl Linnaeus, Christiaan Hendrik Persoon, and Elias Magnus Fries, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or physiology. Advances in molecular genetics have opened the way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. Phylogenetic studies published in the first decade of the 21st century have helped reshape the classification within the fungi kingdom, which is divided into one subkingdom, seven phyla, and ten subphyla.
Etymology
The English word fungus is directly adopted from the Latin fungus (mushroom), used in the writings of Horace and Pliny. This in turn is derived from the Greek word sphongos (σφόγγος 'sponge'), which refers to the macroscopic structures and morphology of mushrooms and molds; the root is also used in other languages, such as the German Schwamm ('sponge') and Schimmel ('mold').
The word mycology is derived from the Greek mykes (μύκης 'mushroom') and logos (λόγος 'discourse'). It denotes the scientific study of fungi. The Latin adjectival form of "mycology" (mycologicæ) appeared as early as 1796 in a book on the subject by Christiaan Hendrik Persoon. The word appeared in English as early as 1824 in a book by Robert Kaye Greville. In 1836 the English naturalist Miles Joseph Berkeley's publication The English Flora of Sir James Edward Smith, Vol. 5. also refers to mycology as the study of fungi.
A group of all the fungi present in a particular region is known as mycobiota (plural noun, no singular). The term mycota is often used for this purpose, but many authors use it as a synonym of Fungi. The word funga has been proposed as a less ambiguous term morphologically similar to fauna and flora. The Species Survival Commission (SSC) of the International Union for Conservation of Nature (IUCN) in August 2021 asked that the phrase fauna and flora be replaced by fauna, flora, and funga.
Characteristics
Fungal hyphae cells
Hyphal wall
Septum
Mitochondrion
Vacuole
Ergosterol crystal
Ribosome
Nucleus
Endoplasmic reticulum
Lipid body
Plasma membrane
Spitzenkörper
Golgi apparatus
Fungal cell cycle showing Dikaryons typical of Higher Fungi
Before the introduction of molecular methods for phylogenetic analysis, taxonomists considered fungi to be members of the plant kingdom because of similarities in lifestyle: both fungi and plants are mainly immobile, and have similarities in general morphology and growth habitat. Although inaccurate, the common misconception that fungi are plants persists among the general public due to their historical classification, as well as several similarities. Like plants, fungi often grow in soil and, in the case of mushrooms, form conspicuous fruit bodies, which sometimes resemble plants such as mosses. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have diverged around one billion years ago (around the start of the Neoproterozoic Era). Some morphological, biochemical, and genetic features are shared with other organisms, while others are unique to the fungi, clearly separating them from the other kingdoms:
With other eukaryotes: Fungal cells contain membrane-bound nuclei with chromosomes that contain DNA with noncoding regions called introns and coding regions called exons. Fungi have membrane-bound cytoplasmic organelles such as mitochondria, sterol-containing membranes, and ribosomes of the 80S type. They have a characteristic range of soluble carbohydrates and storage compounds, including sugar alcohols (e.g., mannitol), disaccharides, (e.g., trehalose), and polysaccharides (e.g., glycogen, which is also found in animals).
With animals: Fungi lack chloroplasts and are heterotrophic organisms and so require preformed organic compounds as energy sources.
With plants: Fungi have a cell wall and vacuoles. They reproduce by both sexual and asexual means, and like basal plant groups (such as ferns and mosses) produce spores. Similar to mosses and algae, fungi typically have haploid nuclei.
With euglenoids and bacteria: Higher fungi, euglenoids, and some bacteria produce the amino acid L-lysine in specific biosynthesis steps, called the α-aminoadipate pathway.
The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called hyphae, which may contain multiple nuclei and extend by growing at their tips. Each tip contains a set of aggregated vesicles—cellular structures consisting of proteins, lipids, and other organic molecules—called the Spitzenkörper. Both fungi and oomycetes grow as filamentous hyphal cells. In contrast, similar-looking organisms, such as filamentous green algae, grow by repeated cell division within a chain of cells. There are also single-celled fungi (yeasts) that do not form hyphae, and some fungi have both hyphal and yeast forms.
In common with some plant and animal species, more than one hundred fungal species display bioluminescence.
Unique features:
Some species grow as unicellular yeasts that reproduce by budding or fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to environmental conditions.
The fungal cell wall is made of a chitin-glucan complex; while glucans are also found in plants and chitin in the exoskeleton of arthropods, fungi are the only organisms that combine these two structural molecules in their cell wall. Unlike those of plants and oomycetes, fungal cell walls do not contain cellulose.
A whitish fan or funnel-shaped mushroom growing at the base of a tree.
Omphalotus nidiformis, a bioluminescent mushroom
Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the xylem and phloem in many plants. To overcome this limitation, some fungi, such as Armillaria, form rhizomorphs, which resemble and perform functions similar to the roots of plants. As eukaryotes, fungi possess a biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate as chemical building blocks. Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure that fungi and animals do not have. Fungi produce several secondary metabolites that are similar or identical in structure to those made by plants. Many of the plant and fungal enzymes that make these compounds differ from each other in sequence and other characteristics, which indicates separate origins and convergent evolution of these enzymes in the fungi and plants.
Diversity
Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as deserts or areas with high salt concentrations or ionizing radiation, as well as in deep sea sediments. Some can survive the intense UV and cosmic radiation encountered during space travel. Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the chytrid fungi Batrachochytrium dendrobatidis and B. salamandrivorans, parasites that have been responsible for a worldwide decline in amphibian populations. These organisms spend part of their life cycle as a motile zoospore, enabling them to propel itself through water and enter their amphibian host. Other examples of aquatic fungi include those living in hydrothermal areas of the ocean.
As of 2020, around 148,000 species of fungi have been described by taxonomists, but the global biodiversity of the fungus kingdom is not fully understood. A 2017 estimate suggests there may be between 2.2 and 3.8 million species The number of new fungi species discovered yearly has increased from 1,000 to 1,500 per year about 10 years ago, to about 2000 with a peak of more than 2,500 species in 2016. In the year 2019, 1882 new species of fungi were described, and it was estimated that more than 90% of fungi remain unknown The following year, 2905 new species were described—the highest annual record of new fungus names. In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on morphological characteristics, such as the size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy. Species may also be distinguished by their biochemical and physiological characteristics, such as their ability to metabolize certain biochemicals, or their reaction to chemical tests. The biological species concept discriminates species based on their ability to mate. The application of molecular tools, such as DNA sequencing and phylogenetic analysis, to study diversity has greatly enhanced the resolution and added robustness to estimates of genetic diversity within various taxonomic groups.
Mycology
Mycology is the branch of biology concerned with the systematic study of fungi, including their genetic and biochemical properties, their taxonomy, and their use to humans as a source of medicine, food, and psychotropic substances consumed for religious purposes, as well as their dangers, such as poisoning or infection. The field of phytopathology, the study of plant diseases, is closely related because many plant pathogens are fungi.
The use of fungi by humans dates back to prehistory; Ötzi the Iceman, a well-preserved mummy of a 5,300-year-old Neolithic man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as tinder (Fomes fomentarius), or for medicinal purposes (Piptoporus betulinus). Ancient peoples have used fungi as food sources—often unknowingly—for millennia, in the preparation of leavened bread and fermented juices. Some of the oldest written records contain references to the destruction of crops that were probably caused by pathogenic fungi.
History
Mycology became a systematic science after the development of the microscope in the 17th century. Although fungal spores were first observed by Giambattista della Porta in 1588, the seminal work in the development of mycology is considered to be the publication of Pier Antonio Micheli's 1729 work Nova plantarum genera. Micheli not only observed spores but also showed that, under the proper conditions, they could be induced into growing into the same species of fungi from which they originated. Extending the use of the binomial system of nomenclature introduced by Carl Linnaeus in his Species plantarum (1753), the Dutch Christiaan Hendrik Persoon (1761–1836) established the first classification of mushrooms with such skill as to be considered a founder of modern mycology. Later, Elias Magnus Fries (1794–1878) further elaborated the classification of fungi, using spore color and microscopic characteristics, methods still used by taxonomists today. Other notable early contributors to mycology in the 17th–19th and early 20th centuries include Miles Joseph Berkeley, August Carl Joseph Corda, Anton de Bary, the brothers Louis René and Charles Tulasne, Arthur H. R. Buller, Curtis G. Lloyd, and Pier Andrea Saccardo. In the 20th and 21st centuries, advances in biochemistry, genetics, molecular biology, biotechnology, DNA sequencing and phylogenetic analysis has provided new insights into fungal relationships and biodiversity, and has challenged traditional morphology-based groupings in fungal taxonomy.
Morphology
Microscopic structures
Monochrome micrograph showing Penicillium hyphae as long, transparent, tube-like structures a few micrometres across. Conidiophores branch out laterally from the hyphae, terminating in bundles of phialides on which spherical condidiophores are arranged like beads on a string. Septa are faintly visible as dark lines crossing the hyphae.
An environmental isolate of Penicillium
Hypha
Conidiophore
Phialide
Conidia
Septa
Most fungi grow as hyphae, which are cylindrical, thread-like structures 2–10 µm in diameter and up to several centimeters in length. Hyphae grow at their tips (apices); new hyphae are typically formed by emergence of new tips along existing hyphae by a process called branching, or occasionally growing hyphal tips fork, giving rise to two parallel-growing hyphae. Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or anastomosis). These growth processes lead to the development of a mycelium, an interconnected network of hyphae. Hyphae can be either septate or coenocytic. Septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at right angles to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized. Septa have pores that allow cytoplasm, organelles, and sometimes nuclei to pass through; an example is the dolipore septum in fungi of the phylum Basidiomycota. Coenocytic hyphae are in essence multinucleate supercells.
Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include haustoria in plant-parasitic species of most fungal phyla,[63] and arbuscules of several mycorrhizal fungi, which penetrate into the host cells to consume nutrients.
Although fungi are opisthokonts—a grouping of evolutionarily related organisms broadly characterized by a single posterior flagellum—all phyla except for the chytrids have lost their posterior flagella. Fungi are unusual among the eukaryotes in having a cell wall that, in addition to glucans (e.g., β-1,3-glucan) and other typical components, also contains the biopolymer chitin.
Macroscopic structures
Fungal mycelia can become visible to the naked eye, for example, on various surfaces and substrates, such as damp walls and spoiled food, where they are commonly called molds. Mycelia grown on solid agar media in laboratory petri dishes are usually referred to as colonies. These colonies can exhibit growth shapes and colors (due to spores or pigmentation) that can be used as diagnostic features in the identification of species or groups. Some individual fungal colonies can reach extraordinary dimensions and ages as in the case of a clonal colony of Armillaria solidipes, which extends over an area of more than 900 ha (3.5 square miles), with an estimated age of nearly 9,000 years.
The apothecium—a specialized structure important in sexual reproduction in the ascomycetes—is a cup-shaped fruit body that is often macroscopic and holds the hymenium, a layer of tissue containing the spore-bearing cells. The fruit bodies of the basidiomycetes (basidiocarps) and some ascomycetes can sometimes grow very large, and many are well known as mushrooms.
Growth and physiology
Time-lapse photography sequence of a peach becoming progressively discolored and disfigured
Mold growth covering a decaying peach. The frames were taken approximately 12 hours apart over a period of six days.
The growth of fungi as hyphae on or in solid substrates or as single cells in aquatic environments is adapted for the efficient extraction of nutrients, because these growth forms have high surface area to volume ratios. Hyphae are specifically adapted for growth on solid surfaces, and to invade substrates and tissues. They can exert large penetrative mechanical forces; for example, many plant pathogens, including Magnaporthe grisea, form a structure called an appressorium that evolved to puncture plant tissues.[71] The pressure generated by the appressorium, directed against the plant epidermis, can exceed 8 megapascals (1,200 psi).[71] The filamentous fungus Paecilomyces lilacinus uses a similar structure to penetrate the eggs of nematodes.
The mechanical pressure exerted by the appressorium is generated from physiological processes that increase intracellular turgor by producing osmolytes such as glycerol. Adaptations such as these are complemented by hydrolytic enzymes secreted into the environment to digest large organic molecules—such as polysaccharides, proteins, and lipids—into smaller molecules that may then be absorbed as nutrients. The vast majority of filamentous fungi grow in a polar fashion (extending in one direction) by elongation at the tip (apex) of the hypha. Other forms of fungal growth include intercalary extension (longitudinal expansion of hyphal compartments that are below the apex) as in the case of some endophytic fungi, or growth by volume expansion during the development of mushroom stipes and other large organs. Growth of fungi as multicellular structures consisting of somatic and reproductive cells—a feature independently evolved in animals and plants—has several functions, including the development of fruit bodies for dissemination of sexual spores (see above) and biofilms for substrate colonization and intercellular communication.
Fungi are traditionally considered heterotrophs, organisms that rely solely on carbon fixed by other organisms for metabolism. Fungi have evolved a high degree of metabolic versatility that allows them to use a diverse range of organic substrates for growth, including simple compounds such as nitrate, ammonia, acetate, or ethanol. In some species the pigment melanin may play a role in extracting energy from ionizing radiation, such as gamma radiation. This form of "radiotrophic" growth has been described for only a few species, the effects on growth rates are small, and the underlying biophysical and biochemical processes are not well known. This process might bear similarity to CO2 fixation via visible light, but instead uses ionizing radiation as a source of energy.
Reproduction
Two thickly stemmed brownish mushrooms with scales on the upper surface, growing out of a tree trunk
Polyporus squamosus
Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms. It is estimated that a third of all fungi reproduce using more than one method of propagation; for example, reproduction may occur in two well-differentiated stages within the life cycle of a species, the teleomorph (sexual reproduction) and the anamorph (asexual reproduction). Environmental conditions trigger genetically determined developmental states that lead to the creation of specialized structures for sexual or asexual reproduction. These structures aid reproduction by efficiently dispersing spores or spore-containing propagules.
Asexual reproduction
Asexual reproduction occurs via vegetative spores (conidia) or through mycelial fragmentation. Mycelial fragmentation occurs when a fungal mycelium separates into pieces, and each component grows into a separate mycelium. Mycelial fragmentation and vegetative spores maintain clonal populations adapted to a specific niche, and allow more rapid dispersal than sexual reproduction. The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or Deuteromycota comprise all the species that lack an observable sexual cycle. Deuteromycota (alternatively known as Deuteromycetes, conidial fungi, or mitosporic fungi) is not an accepted taxonomic clade and is now taken to mean simply fungi that lack a known sexual stage.
Sexual reproduction
See also: Mating in fungi and Sexual selection in fungi
Sexual reproduction with meiosis has been directly observed in all fungal phyla except Glomeromycota (genetic analysis suggests meiosis in Glomeromycota as well). It differs in many aspects from sexual reproduction in animals or plants. Differences also exist between fungal groups and can be used to discriminate species by morphological differences in sexual structures and reproductive strategies. Mating experiments between fungal isolates may identify species on the basis of biological species concepts. The major fungal groupings have initially been delineated based on the morphology of their sexual structures and spores; for example, the spore-containing structures, asci and basidia, can be used in the identification of ascomycetes and basidiomycetes, respectively. Fungi employ two mating systems: heterothallic species allow mating only between individuals of the opposite mating type, whereas homothallic species can mate, and sexually reproduce, with any other individual or itself.
Most fungi have both a haploid and a diploid stage in their life cycles. In sexually reproducing fungi, compatible individuals may combine by fusing their hyphae together into an interconnected network; this process, anastomosis, is required for the initiation of the sexual cycle. Many ascomycetes and basidiomycetes go through a dikaryotic stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see heterokaryosis).
In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing tissue layer) form a characteristic hook (crozier) at the hyphal septum. During cell division, the formation of the hook ensures proper distribution of the newly divided nuclei into the apical and basal hyphal compartments. An ascus (plural asci) is then formed, in which karyogamy (nuclear fusion) occurs. Asci are embedded in an ascocarp, or fruiting body. Karyogamy in the asci is followed immediately by meiosis and the production of ascospores. After dispersal, the ascospores may germinate and form a new haploid mycelium.
Sexual reproduction in basidiomycetes is similar to that of the ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic mycelium. However, the dikaryotic phase is more extensive in the basidiomycetes, often also present in the vegetatively growing mycelium. A specialized anatomical structure, called a clamp connection, is formed at each hyphal septum. As with the structurally similar hook in the ascomycetes, the clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two genetically different nuclei in each hyphal compartment. A basidiocarp is formed in which club-like structures known as basidia generate haploid basidiospores after karyogamy and meiosis. The most commonly known basidiocarps are mushrooms, but they may also take other forms (see Morphology section).
In fungi formerly classified as Zygomycota, haploid hyphae of two individuals fuse, forming a gametangium, a specialized cell structure that becomes a fertile gamete-producing cell. The gametangium develops into a zygospore, a thick-walled spore formed by the union of gametes. When the zygospore germinates, it undergoes meiosis, generating new haploid hyphae, which may then form asexual sporangiospores. These sporangiospores allow the fungus to rapidly disperse and germinate into new genetically identical haploid fungal mycelia.
Spore dispersal
The spores of most of the researched species of fungi are transported by wind. Such species often produce dry or hydrophobic spores that do not absorb water and are readily scattered by raindrops, for example. In other species, both asexual and sexual spores or sporangiospores are often actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of the spores from the reproductive structures as well as traveling through the air over long distances.
Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as hydrophobins), enable efficient spore ejection. For example, the structure of the spore-bearing cells in some ascomycete species is such that the buildup of substances affecting cell volume and fluid balance enables the explosive discharge of spores into the air. The forcible discharge of single spores termed ballistospores involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g; the net result is that the spore is ejected 0.01–0.02 cm, sufficient distance for it to fall through the gills or pores into the air below. Other fungi, like the puffballs, rely on alternative mechanisms for spore release, such as external mechanical forces. The hydnoid fungi (tooth fungi) produce spores on pendant, tooth-like or spine-like projections. The bird's nest fungi use the force of falling water drops to liberate the spores from cup-shaped fruiting bodies. Another strategy is seen in the stinkhorns, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.
Homothallism
In homothallic sexual reproduction, two haploid nuclei derived from the same individual fuse to form a zygote that can then undergo meiosis. Homothallic fungi include species with an Aspergillus-like asexual stage (anamorphs) occurring in numerous different genera, several species of the ascomycete genus Cochliobolus, and the ascomycete Pneumocystis jirovecii. The earliest mode of sexual reproduction among eukaryotes was likely homothallism, that is, self-fertile unisexual reproduction.
Other sexual processes
Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera Penicillium and Aspergillus, may exchange genetic material via parasexual processes, initiated by anastomosis between hyphae and plasmogamy of fungal cells. The frequency and relative importance of parasexual events is unclear and may be lower than other sexual processes. It is known to play a role in intraspecific hybridization and is likely required for hybridization between species, which has been associated with major events in fungal evolution.
Evolution
In contrast to plants and animals, the early fossil record of the fungi is meager. Factors that likely contribute to the under-representation of fungal species among fossils include the nature of fungal fruiting bodies, which are soft, fleshy, and easily degradable tissues and the microscopic dimensions of most fungal structures, which therefore are not readily evident. Fungal fossils are difficult to distinguish from those of other microbes, and are most easily identified when they resemble extant fungi. Often recovered from a permineralized plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with light microscopy or transmission electron microscopy. Researchers study compression fossils by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.
The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some 2,400 million years ago (Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis. Other studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups. The oldest fossilizied mycelium to be identified from its molecular composition is between 715 and 810 million years old. For much of the Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant chytrids in having flagellum-bearing spores. The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including parasitism, saprobism, and the development of mutualistic relationships such as mycorrhiza and lichenization. Studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.
In May 2019, scientists reported the discovery of a fossilized fungus, named Ourasphaira giraldae, in the Canadian Arctic, that may have grown on land a billion years ago, well before plants were living on land. Pyritized fungus-like microfossils preserved in the basal Ediacaran Doushantuo Formation (~635 Ma) have been reported in South China. Earlier, it had been presumed that the fungi colonized the land during the Cambrian (542–488.3 Ma), also long before land plants. Fossilized hyphae and spores recovered from the Ordovician of Wisconsin (460 Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular bryophyte-like plants. Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian and early Devonian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2 Ma), when they occur abundantly in the Rhynie chert, mostly as Zygomycota and Chytridiomycota. At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged, and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).
Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 415 Ma; this date roughly corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie Chert. The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to the mushroom-producing species of the Agaricomycetes). Two amber-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species Archaeomarasmius leggetti) appeared during the late Cretaceous, 90 Ma.
Some time after the Permian–Triassic extinction event (251.4 Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available fossil record for this period. However, the relative proportion of fungal spores relative to spores formed by algal species is difficult to assess, the spike did not appear worldwide, and in many places it did not fall on the Permian–Triassic boundary.
Sixty-five million years ago, immediately after the Cretaceous–Paleogene extinction event that famously killed off most dinosaurs, there was a dramatic increase in evidence of fungi; apparently the death of most plant and animal species led to a huge fungal bloom like "a massive compost heap".
Taxonomy
Although commonly included in botany curricula and textbooks, fungi are more closely related to animals than to plants and are placed with the animals in the monophyletic group of opisthokonts. Analyses using molecular phylogenetics support a monophyletic origin of fungi. The taxonomy of fungi is in a state of constant flux, especially due to research based on DNA comparisons. These current phylogenetic analyses often overturn classifications based on older and sometimes less discriminative methods based on morphological features and biological species concepts obtained from experimental matings.
There is no unique generally accepted system at the higher taxonomic levels and there are frequent name changes at every level, from species upwards. Efforts among researchers are now underway to establish and encourage usage of a unified and more consistent nomenclature. Until relatively recent (2012) changes to the International Code of Nomenclature for algae, fungi and plants, fungal species could also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as Index Fungorum and MycoBank are officially recognized nomenclatural repositories and list current names of fungal species (with cross-references to older synonyms).
The 2007 classification of Kingdom Fungi is the result of a large-scale collaborative research effort involving dozens of mycologists and other scientists working on fungal taxonomy. It recognizes seven phyla, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing subkingdom Dikarya, the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and the beer, wine, and bread yeasts. The accompanying cladogram depicts the major fungal taxa and their relationship to opisthokont and unikont organisms, based on the work of Philippe Silar, "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research" and Tedersoo et al. 2018. The lengths of the branches are not proportional to evolutionary distances.
The major phyla (sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual reproductive structures. As of 2019, nine major lineages have been identified: Opisthosporidia, Chytridiomycota, Neocallimastigomycota, Blastocladiomycota, Zoopagomycotina, Mucoromycota, Glomeromycota, Ascomycota and Basidiomycota.
Phylogenetic analysis has demonstrated that the Microsporidia, unicellular parasites of animals and protists, are fairly recent and highly derived endobiotic fungi (living within the tissue of another species). Previously considered to be "primitive" protozoa, they are now thought to be either a basal branch of the Fungi, or a sister group–each other's closest evolutionary relative.
The Chytridiomycota are commonly known as chytrids. These fungi are distributed worldwide. Chytrids and their close relatives Neocallimastigomycota and Blastocladiomycota (below) are the only fungi with active motility, producing zoospores that are capable of active movement through aqueous phases with a single flagellum, leading early taxonomists to classify them as protists. Molecular phylogenies, inferred from rRNA sequences in ribosomes, suggest that the Chytrids are a basal group divergent from the other fungal phyla, consisting of four major clades with suggestive evidence for paraphyly or possibly polyphyly.
The Blastocladiomycota were previously considered a taxonomic clade within the Chytridiomycota. Molecular data and ultrastructural characteristics, however, place the Blastocladiomycota as a sister clade to the Zygomycota, Glomeromycota, and Dikarya (Ascomycota and Basidiomycota). The blastocladiomycetes are saprotrophs, feeding on decomposing organic matter, and they are parasites of all eukaryotic groups. Unlike their close relatives, the chytrids, most of which exhibit zygotic meiosis, the blastocladiomycetes undergo sporic meiosis.
The Neocallimastigomycota were earlier placed in the phylum Chytridiomycota. Members of this small phylum are anaerobic organisms, living in the digestive system of larger herbivorous mammals and in other terrestrial and aquatic environments enriched in cellulose (e.g., domestic waste landfill sites). They lack mitochondria but contain hydrogenosomes of mitochondrial origin. As in the related chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or polyflagellate.
Microscopic view of a layer of translucent grayish cells, some containing small dark-color spheres
Arbuscular mycorrhiza seen under microscope. Flax root cortical cells containing paired arbuscules.
Cross-section of a cup-shaped structure showing locations of developing meiotic asci (upper edge of cup, left side, arrows pointing to two gray cells containing four and two small circles), sterile hyphae (upper edge of cup, right side, arrows pointing to white cells with a single small circle in them), and mature asci (upper edge of cup, pointing to two gray cells with eight small circles in them)
Diagram of an apothecium (the typical cup-like reproductive structure of Ascomycetes) showing sterile tissues as well as developing and mature asci.
Members of the Glomeromycota form arbuscular mycorrhizae, a form of mutualist symbiosis wherein fungal hyphae invade plant root cells and both species benefit from the resulting increased supply of nutrients. All known Glomeromycota species reproduce asexually. The symbiotic association between the Glomeromycota and plants is ancient, with evidence dating to 400 million years ago. Formerly part of the Zygomycota (commonly known as 'sugar' and 'pin' molds), the Glomeromycota were elevated to phylum status in 2001 and now replace the older phylum Zygomycota. Fungi that were placed in the Zygomycota are now being reassigned to the Glomeromycota, or the subphyla incertae sedis Mucoromycotina, Kickxellomycotina, the Zoopagomycotina and the Entomophthoromycotina. Some well-known examples of fungi formerly in the Zygomycota include black bread mold (Rhizopus stolonifer), and Pilobolus species, capable of ejecting spores several meters through the air. Medically relevant genera include Mucor, Rhizomucor, and Rhizopus.
The Ascomycota, commonly known as sac fungi or ascomycetes, constitute the largest taxonomic group within the Eumycota. These fungi form meiotic spores called ascospores, which are enclosed in a special sac-like structure called an ascus. This phylum includes morels, a few mushrooms and truffles, unicellular yeasts (e.g., of the genera Saccharomyces, Kluyveromyces, Pichia, and Candida), and many filamentous fungi living as saprotrophs, parasites, and mutualistic symbionts (e.g. lichens). Prominent and important genera of filamentous ascomycetes include Aspergillus, Penicillium, Fusarium, and Claviceps. Many ascomycete species have only been observed undergoing asexual reproduction (called anamorphic species), but analysis of molecular data has often been able to identify their closest teleomorphs in the Ascomycota. Because the products of meiosis are retained within the sac-like ascus, ascomycetes have been used for elucidating principles of genetics and heredity (e.g., Neurospora crassa).
Members of the Basidiomycota, commonly known as the club fungi or basidiomycetes, produce meiospores called basidiospores on club-like stalks called basidia. Most common mushrooms belong to this group, as well as rust and smut fungi, which are major pathogens of grains. Other important basidiomycetes include the maize pathogen Ustilago maydis, human commensal species of the genus Malassezia, and the opportunistic human pathogen, Cryptococcus neoformans.
Fungus-like organisms
Because of similarities in morphology and lifestyle, the slime molds (mycetozoans, plasmodiophorids, acrasids, Fonticula and labyrinthulids, now in Amoebozoa, Rhizaria, Excavata, Opisthokonta and Stramenopiles, respectively), water molds (oomycetes) and hyphochytrids (both Stramenopiles) were formerly classified in the kingdom Fungi, in groups like Mastigomycotina, Gymnomycota and Phycomycetes. The slime molds were studied also as protozoans, leading to an ambiregnal, duplicated taxonomy.
Unlike true fungi, the cell walls of oomycetes contain cellulose and lack chitin. Hyphochytrids have both chitin and cellulose. Slime molds lack a cell wall during the assimilative phase (except labyrinthulids, which have a wall of scales), and take in nutrients by ingestion (phagocytosis, except labyrinthulids) rather than absorption (osmotrophy, as fungi, labyrinthulids, oomycetes and hyphochytrids). Neither water molds nor slime molds are closely related to the true fungi, and, therefore, taxonomists no longer group them in the kingdom Fungi. Nonetheless, studies of the oomycetes and myxomycetes are still often included in mycology textbooks and primary research literature.
The Eccrinales and Amoebidiales are opisthokont protists, previously thought to be zygomycete fungi. Other groups now in Opisthokonta (e.g., Corallochytrium, Ichthyosporea) were also at given time classified as fungi. The genus Blastocystis, now in Stramenopiles, was originally classified as a yeast. Ellobiopsis, now in Alveolata, was considered a chytrid. The bacteria were also included in fungi in some classifications, as the group Schizomycetes.
The Rozellida clade, including the "ex-chytrid" Rozella, is a genetically disparate group known mostly from environmental DNA sequences that is a sister group to fungi. Members of the group that have been isolated lack the chitinous cell wall that is characteristic of fungi. Alternatively, Rozella can be classified as a basal fungal group.
The nucleariids may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom. Many Actinomycetales (Actinomycetota), a group with many filamentous bacteria, were also long believed to be fungi.
Ecology
Although often inconspicuous, fungi occur in every environment on Earth and play very important roles in most ecosystems. Along with bacteria, fungi are the major decomposers in most terrestrial (and some aquatic) ecosystems, and therefore play a critical role in biogeochemical cycles and in many food webs. As decomposers, they play an essential role in nutrient cycling, especially as saprotrophs and symbionts, degrading organic matter to inorganic molecules, which can then re-enter anabolic metabolic pathways in plants or other organisms.
Symbiosis
Many fungi have important symbiotic relationships with organisms from most if not all kingdoms. These interactions can be mutualistic or antagonistic in nature, or in the case of commensal fungi are of no apparent benefit or detriment to the host.
With plants
Mycorrhizal symbiosis between plants and fungi is one of the most well-known plant–fungus associations and is of significant importance for plant growth and persistence in many ecosystems; over 90% of all plant species engage in mycorrhizal relationships with fungi and are dependent upon this relationship for survival.
A microscopic view of blue-stained cells, some with dark wavy lines in them
The dark filaments are hyphae of the endophytic fungus Epichloë coenophiala in the intercellular spaces of tall fescue leaf sheath tissue
The mycorrhizal symbiosis is ancient, dating back to at least 400 million years. It often increases the plant's uptake of inorganic compounds, such as nitrate and phosphate from soils having low concentrations of these key plant nutrients. The fungal partners may also mediate plant-to-plant transfer of carbohydrates and other nutrients. Such mycorrhizal communities are called "common mycorrhizal networks". A special case of mycorrhiza is myco-heterotrophy, whereby the plant parasitizes the fungus, obtaining all of its nutrients from its fungal symbiont. Some fungal species inhabit the tissues inside roots, stems, and leaves, in which case they are called endophytes. Similar to mycorrhiza, endophytic colonization by fungi may benefit both symbionts; for example, endophytes of grasses impart to their host increased resistance to herbivores and other environmental stresses and receive food and shelter from the plant in return.
With algae and cyanobacteria
A green, leaf-like structure attached to a tree, with a pattern of ridges and depression on the bottom surface
The lichen Lobaria pulmonaria, a symbiosis of fungal, algal, and cyanobacterial species
Lichens are a symbiotic relationship between fungi and photosynthetic algae or cyanobacteria. The photosynthetic partner in the relationship is referred to in lichen terminology as a "photobiont". The fungal part of the relationship is composed mostly of various species of ascomycetes and a few basidiomycetes. Lichens occur in every ecosystem on all continents, play a key role in soil formation and the initiation of biological succession, and are prominent in some extreme environments, including polar, alpine, and semiarid desert regions. They are able to grow on inhospitable surfaces, including bare soil, rocks, tree bark, wood, shells, barnacles and leaves. As in mycorrhizas, the photobiont provides sugars and other carbohydrates via photosynthesis to the fungus, while the fungus provides minerals and water to the photobiont. The functions of both symbiotic organisms are so closely intertwined that they function almost as a single organism; in most cases the resulting organism differs greatly from the individual components. Lichenization is a common mode of nutrition for fungi; around 27% of known fungi—more than 19,400 species—are lichenized. Characteristics common to most lichens include obtaining organic carbon by photosynthesis, slow growth, small size, long life, long-lasting (seasonal) vegetative reproductive structures, mineral nutrition obtained largely from airborne sources, and greater tolerance of desiccation than most other photosynthetic organisms in the same habitat.
With insects
Many insects also engage in mutualistic relationships with fungi. Several groups of ants cultivate fungi in the order Chaetothyriales for several purposes: as a food source, as a structural component of their nests, and as a part of an ant/plant symbiosis in the domatia (tiny chambers in plants that house arthropods). Ambrosia beetles cultivate various species of fungi in the bark of trees that they infest. Likewise, females of several wood wasp species (genus Sirex) inject their eggs together with spores of the wood-rotting fungus Amylostereum areolatum into the sapwood of pine trees; the growth of the fungus provides ideal nutritional conditions for the development of the wasp larvae. At least one species of stingless bee has a relationship with a fungus in the genus Monascus, where the larvae consume and depend on fungus transferred from old to new nests. Termites on the African savannah are also known to cultivate fungi, and yeasts of the genera Candida and Lachancea inhabit the gut of a wide range of insects, including neuropterans, beetles, and cockroaches; it is not known whether these fungi benefit their hosts. Fungi growing in dead wood are essential for xylophagous insects (e.g. woodboring beetles). They deliver nutrients needed by xylophages to nutritionally scarce dead wood. Thanks to this nutritional enrichment the larvae of the woodboring insect is able to grow and develop to adulthood. The larvae of many families of fungicolous flies, particularly those within the superfamily Sciaroidea such as the Mycetophilidae and some Keroplatidae feed on fungal fruiting bodies and sterile mycorrhizae.
A thin brown stick positioned horizontally with roughly two dozen clustered orange-red leaves originating from a single point in the middle of the stick. These orange leaves are three to four times larger than the few other green leaves growing out of the stick, and are covered on the lower leaf surface with hundreds of tiny bumps. The background shows the green leaves and branches of neighboring shrubs.
The plant pathogen Puccinia magellanicum (calafate rust) causes the defect known as witch's broom, seen here on a barberry shrub in Chile.
Gram stain of Candida albicans from a vaginal swab from a woman with candidiasis, showing hyphae, and chlamydospores, which are 2–4 µm in diameter.
Many fungi are parasites on plants, animals (including humans), and other fungi. Serious pathogens of many cultivated plants causing extensive damage and losses to agriculture and forestry include the rice blast fungus Magnaporthe oryzae, tree pathogens such as Ophiostoma ulmi and Ophiostoma novo-ulmi causing Dutch elm disease, Cryphonectria parasitica responsible for chestnut blight, and Phymatotrichopsis omnivora causing Texas Root Rot, and plant pathogens in the genera Fusarium, Ustilago, Alternaria, and Cochliobolus. Some carnivorous fungi, like Paecilomyces lilacinus, are predators of nematodes, which they capture using an array of specialized structures such as constricting rings or adhesive nets. Many fungi that are plant pathogens, such as Magnaporthe oryzae, can switch from being biotrophic (parasitic on living plants) to being necrotrophic (feeding on the dead tissues of plants they have killed). This same principle is applied to fungi-feeding parasites, including Asterotremella albida, which feeds on the fruit bodies of other fungi both while they are living and after they are dead.
Some fungi can cause serious diseases in humans, several of which may be fatal if untreated. These include aspergillosis, candidiasis, coccidioidomycosis, cryptococcosis, histoplasmosis, mycetomas, and paracoccidioidomycosis. Furthermore, persons with immuno-deficiencies are particularly susceptible to disease by genera such as Aspergillus, Candida, Cryptoccocus, Histoplasma, and Pneumocystis. Other fungi can attack eyes, nails, hair, and especially skin, the so-called dermatophytic and keratinophilic fungi, and cause local infections such as ringworm and athlete's foot. Fungal spores are also a cause of allergies, and fungi from different taxonomic groups can evoke allergic reactions.
As targets of mycoparasites
Organisms that parasitize fungi are known as mycoparasitic organisms. About 300 species of fungi and fungus-like organisms, belonging to 13 classes and 113 genera, are used as biocontrol agents against plant fungal diseases. Fungi can also act as mycoparasites or antagonists of other fungi, such as Hypomyces chrysospermus, which grows on bolete mushrooms. Fungi can also become the target of infection by mycoviruses.
Communication
Main article: Mycorrhizal networks
There appears to be electrical communication between fungi in word-like components according to spiking characteristics.
Possible impact on climate
According to a study published in the academic journal Current Biology, fungi can soak from the atmosphere around 36% of global fossil fuel greenhouse gas emissions.
Mycotoxins
(6aR,9R)-N-((2R,5S,10aS,10bS)-5-benzyl-10b-hydroxy-2-methyl-3,6-dioxooctahydro-2H-oxazolo[3,2-a] pyrrolo[2,1-c]pyrazin-2-yl)-7-methyl-4,6,6a,7,8,9-hexahydroindolo[4,3-fg] quinoline-9-carboxamide
Ergotamine, a major mycotoxin produced by Claviceps species, which if ingested can cause gangrene, convulsions, and hallucinations
Many fungi produce biologically active compounds, several of which are toxic to animals or plants and are therefore called mycotoxins. Of particular relevance to humans are mycotoxins produced by molds causing food spoilage, and poisonous mushrooms (see above). Particularly infamous are the lethal amatoxins in some Amanita mushrooms, and ergot alkaloids, which have a long history of causing serious epidemics of ergotism (St Anthony's Fire) in people consuming rye or related cereals contaminated with sclerotia of the ergot fungus, Claviceps purpurea. Other notable mycotoxins include the aflatoxins, which are insidious liver toxins and highly carcinogenic metabolites produced by certain Aspergillus species often growing in or on grains and nuts consumed by humans, ochratoxins, patulin, and trichothecenes (e.g., T-2 mycotoxin) and fumonisins, which have significant impact on human food supplies or animal livestock.
Mycotoxins are secondary metabolites (or natural products), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi. Mycotoxins may provide fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption (fungivory). Many fungal secondary metabolites (or derivatives) are used medically, as described under Human use below.
Pathogenic mechanisms
Ustilago maydis is a pathogenic plant fungus that causes smut disease in maize and teosinte. Plants have evolved efficient defense systems against pathogenic microbes such as U. maydis. A rapid defense reaction after pathogen attack is the oxidative burst where the plant produces reactive oxygen species at the site of the attempted invasion. U. maydis can respond to the oxidative burst with an oxidative stress response, regulated by the gene YAP1. The response protects U. maydis from the host defense, and is necessary for the pathogen's virulence. Furthermore, U. maydis has a well-established recombinational DNA repair system which acts during mitosis and meiosis. The system may assist the pathogen in surviving DNA damage arising from the host plant's oxidative defensive response to infection.
Cryptococcus neoformans is an encapsulated yeast that can live in both plants and animals. C. neoformans usually infects the lungs, where it is phagocytosed by alveolar macrophages. Some C. neoformans can survive inside macrophages, which appears to be the basis for latency, disseminated disease, and resistance to antifungal agents. One mechanism by which C. neoformans survives the hostile macrophage environment is by up-regulating the expression of genes involved in the oxidative stress response. Another mechanism involves meiosis. The majority of C. neoformans are mating "type a". Filaments of mating "type a" ordinarily have haploid nuclei, but they can become diploid (perhaps by endoduplication or by stimulated nuclear fusion) to form blastospores. The diploid nuclei of blastospores can undergo meiosis, including recombination, to form haploid basidiospores that can be dispersed. This process is referred to as monokaryotic fruiting. This process requires a gene called DMC1, which is a conserved homologue of genes recA in bacteria and RAD51 in eukaryotes, that mediates homologous chromosome pairing during meiosis and repair of DNA double-strand breaks. Thus, C. neoformans can undergo a meiosis, monokaryotic fruiting, that promotes recombinational repair in the oxidative, DNA damaging environment of the host macrophage, and the repair capability may contribute to its virulence.
Human use
See also: Human interactions with fungi
Microscopic view of five spherical structures; one of the spheres is considerably smaller than the rest and attached to one of the larger spheres
Saccharomyces cerevisiae cells shown with DIC microscopy
The human use of fungi for food preparation or preservation and other purposes is extensive and has a long history. Mushroom farming and mushroom gathering are large industries in many countries. The study of the historical uses and sociological impact of fungi is known as ethnomycology. Because of the capacity of this group to produce an enormous range of natural products with antimicrobial or other biological activities, many species have long been used or are being developed for industrial production of antibiotics, vitamins, and anti-cancer and cholesterol-lowering drugs. Methods have been developed for genetic engineering of fungi, enabling metabolic engineering of fungal species. For example, genetic modification of yeast species—which are easy to grow at fast rates in large fermentation vessels—has opened up ways of pharmaceutical production that are potentially more efficient than production by the original source organisms. Fungi-based industries are sometimes considered to be a major part of a growing bioeconomy, with applications under research and development including use for textiles, meat substitution and general fungal biotechnology.
Therapeutic uses
Modern chemotherapeutics
Many species produce metabolites that are major sources of pharmacologically active drugs.
Antibiotics
Particularly important are the antibiotics, including the penicillins, a structurally related group of β-lactam antibiotics that are synthesized from small peptides. Although naturally occurring penicillins such as penicillin G (produced by Penicillium chrysogenum) have a relatively narrow spectrum of biological activity, a wide range of other penicillins can be produced by chemical modification of the natural penicillins. Modern penicillins are semisynthetic compounds, obtained initially from fermentation cultures, but then structurally altered for specific desirable properties. Other antibiotics produced by fungi include: ciclosporin, commonly used as an immunosuppressant during transplant surgery; and fusidic acid, used to help control infection from methicillin-resistant Staphylococcus aureus bacteria. Widespread use of antibiotics for the treatment of bacterial diseases, such as tuberculosis, syphilis, leprosy, and others began in the early 20th century and continues to date. In nature, antibiotics of fungal or bacterial origin appear to play a dual role: at high concentrations they act as chemical defense against competition with other microorganisms in species-rich environments, such as the rhizosphere, and at low concentrations as quorum-sensing molecules for intra- or interspecies signaling.
Other
Other drugs produced by fungi include griseofulvin isolated from Penicillium griseofulvum, used to treat fungal infections, and statins (HMG-CoA reductase inhibitors), used to inhibit cholesterol synthesis. Examples of statins found in fungi include mevastatin from Penicillium citrinum and lovastatin from Aspergillus terreus and the oyster mushroom. Psilocybin from fungi is investigated for therapeutic use and appears to cause global increases in brain network integration. Fungi produce compounds that inhibit viruses and cancer cells. Specific metabolites, such as polysaccharide-K, ergotamine, and β-lactam antibiotics, are routinely used in clinical medicine. The shiitake mushroom is a source of lentinan, a clinical drug approved for use in cancer treatments in several countries, including Japan. In Europe and Japan, polysaccharide-K (brand name Krestin), a chemical derived from Trametes versicolor, is an approved adjuvant for cancer therapy.
Traditional medicine
Upper surface view of a kidney-shaped fungus, brownish-red with a lighter yellow-brown margin, and a somewhat varnished or shiny appearance
Two dried yellow-orange caterpillars, one with a curly grayish fungus growing out of one of its ends. The grayish fungus is roughly equal to or slightly greater in length than the caterpillar, and tapers in thickness to a narrow end.
The fungi Ganoderma lucidum (left) and Ophiocordyceps sinensis (right) are used in traditional medicine practices
Certain mushrooms are used as supposed therapeutics in folk medicine practices, such as traditional Chinese medicine. Mushrooms with a history of such use include Agaricus subrufescens, Ganoderma lucidum, and Ophiocordyceps sinensis.
Cultured foods
Baker's yeast or Saccharomyces cerevisiae, a unicellular fungus, is used to make bread and other wheat-based products, such as pizza dough and dumplings. Yeast species of the genus Saccharomyces are also used to produce alcoholic beverages through fermentation. Shoyu koji mold (Aspergillus oryzae) is an essential ingredient in brewing Shoyu (soy sauce) and sake, and the preparation of miso while Rhizopus species are used for making tempeh. Several of these fungi are domesticated species that were bred or selected according to their capacity to ferment food without producing harmful mycotoxins (see below), which are produced by very closely related Aspergilli. Quorn, a meat substitute, is made from Fusarium venenatum.
Last week-end, we had lunch by the sea, where some people enjoyed the lovely autumnal Mediterranean sunshine.
This week, back home, pijama day instead -wind cold and rain outside.
Tarragona, October 2013.
Do u know the difference between two giants mobile sector, Samsung galaxy S4 and iPhone 5? Lots of people say iPhone 5 is a good phone but Don't ignore Samsung galaxy S4... Latest new mobiles our blog for more info..
Sovage Denim can change your energy! Sovage can make the difference in your day. Sovage Denim can brighten your day with colors to inspire you, and a fit that feels like the skin you were born in. Sovage Denim lifts your spirit, your soul; expresses your freedom! Sovage is life! The denim was born in the fight for freedom. Every part of the denim is snapshot of the culture that symbolizes freedom.
The silk lining depicts the designs that were given to Toussaint by the angels and saints in his dreams and visions that would facilitate victory for his people. These symbols were a secret code to beat the enemy. The colors of the stitching and buttons symbolize the colors of the spirits, sent by God, to help Toussaint win the war for freedom. The buttons symbolize the royal crown Toussaint L'Ouverture earned after living his life as a slave. When you see the different washes, think of the tears that have been cried and the rains that have washed them away.
Made of the highest quality fabrics and with great attention to detail, you will never find another pair of pants like Sovage Denim. This LA-based, high-end designer denim boasts the very best fit in jeans. A fit for all styles, Sovage Denim has something for everyone: vintage to flashy - conservative to sexy.
Celebrities who Rock Sovage Denim includes Aaron Carter, Angel McCord, Bridgetta Tomarchio, Ellen Hollman, Jayde Nicole, Karina Smirnoff, Stephanie Pratt and Tami Farrell
For news and updates on Sovage Denim go to www.SovageDenim.com
Submitted by jati.