title: Dung beetle

text: Dung beetles play a role in agriculture and tropical forests. By burying and consuming dung, they improve nutrient recycling and soil structure. Dung beetles have been further shown to improve soil conditions and plant growth on rehabilitated coal mines in South Africa. They are also important for the dispersal of seeds present in animals' dung, influencing seed burial and seedling recruitment in tropical forests. They can protect livestock, such as cattle, by removing the dung which, if left, could provide habitat for pests such as flies. Therefore, many countries have introduced the creatures for the benefit of animal husbandry. The American Institute of Biological Sciences reports that dung beetles save the United States cattle industry an estimated US$380 million annually through burying above-ground livestock feces.


title: Dung beetle

text: In Australia, the Commonwealth Scientific and Industrial Research Organisation (CSIRO) commissioned the Australian Dung Beetle Project (1965–1985) which, led by George Bornemissza, sought to introduce species of dung beetles from South Africa and Europe. The successful introduction of 23 species was made, most notably "Digitonthophagus gazella" and "Euoniticellus intermedius", which has resulted in improvement of the quality and fertility of Australian cattle pastures, along with a reduction in the population of pestilent Australian bush flies by around 90%. In 1995 it was reported that dung beetles were being trialled in the Sydney beach suburb of Curl Curl to deal with dog droppings.


title: Dung beetle

text: An application has been made by Landcare Research to import up to 11 species of dung beetle into New Zealand. As well as improving pasture soils the Dung Beetle Release Strategy Group say that it would result in a reduction in emissions of nitrous oxide (a greenhouse gas) from agriculture. There is, however, strong opposition from some at the University of Auckland, and a few others, based on the risks of the dung beetles acting as vectors of disease. There are public health researchers at the University of Auckland who agree with the current EPA risk assessment and indeed there are several Landcare programmes in Australia that involve schoolchildren collecting dung beetles.


title: Dung beetle

text: The African dung beetle ("D. gazella") was introduced in several locations in North and South America and has been spreading its distribution to other regions by natural dispersal and accidental transportation, and is now probably naturalized in most countries between México and Argentina. The exotic species might be useful for controlling diseases of livestock in commercial areas, and might displace native species in modified landscapes; however, data is not conclusive about its effect on native species in natural environments and further monitoring is required.


title: Dung beetle

text: Like many other insects, (dried) dung beetle, called "qiāngláng" (蜣蜋) in Chinese, is used in Chinese herbal medicine. It is recorded in the "Insect section" (蟲部) of the "Compendium of Materia Medica", where it is recommended for the cure of 10 diseases.


title: Dung beetle

text: In Isan, Northeastern Thailand, the local people eat many different kinds of insects, including the dung beetle. There is an Isan song กุดจี่หายไปใหน "Where Did the Dung Beetle Go", which relates the replacement of water buffalo with the "metal" buffalo, which does not provide the dung needed for the dung beetle and has led to the increasing rarity of the dung beetle in the agricultural region.


title: Dung beetle

text: The Mediterranean dung beetle ("Bubas bison") has been used in conjunction with biochar stock fodder to reduce emissions of nitrous oxide and carbon dioxide, which are both greenhouse gases. The beetles work the biochar-enriched dung into the soil without the use of machines.


title: Dung beetle

text: Some dung beetles are used as food in South East Asia and a variety of dung beetle species have been used therapeutically (and are still being used in traditionally living societies) in potions and folk medicines to treat a number of illnesses and disorders.


title: Dung beetle

text: Several species of the dung beetle, most notably the species "Scarabaeus sacer" (often referred to as "the sacred scarab"), enjoyed a sacred status among the ancient Egyptians.


title: Dung beetle

text: Egyptian hieroglyphic script uses the image of the beetle to represent a triliteral phonetic that Egyptologists transliterate as "xpr" or "ḫpr" and translate as "to come into being", "to become" or "to transform". The derivative term "xprw" or "ḫpr(w)" is variously translated as "form", "transformation", "happening", "mode of being" or "what has come into being", depending on the context. It may have existential, fictional, or ontologic significance.


title: Dung beetle

text: The scarab was linked to Khepri ("he who has come into being"), the god of the rising sun. The ancients believed that the dung beetle was only male-sexed, and reproduced by depositing semen into a dung ball. The supposed self-creation of the beetle resembles that of Khepri, who creates himself out of nothing. Moreover, the dung ball rolled by a dung beetle resembles the sun. Plutarch wrote:


title: Dung beetle

text: The ancient Egyptians believed that Khepri renewed the sun every day before rolling it above the horizon, then carried it through the other world after sunset, only to renew it, again, the next day. Some New Kingdom royal tombs exhibit a threefold image of the sun god, with the beetle as symbol of the morning sun. The astronomical ceiling in the tomb of Ramses VI portrays the nightly "death" and "rebirth" of the sun as being swallowed by Nut, goddess of the sky, and re-emerging from her womb as Khepri.


title: Dung beetle

text: The image of the scarab, conveying ideas of transformation, renewal, and resurrection, is ubiquitous in ancient Egyptian religious and funerary art.


title: Dung beetle

text: Excavations of ancient Egyptian sites have yielded images of the scarab in bone, ivory, stone, Egyptian faience, and precious metals, dating from the Sixth Dynasty and up to the period of Roman rule. They are generally small, bored to allow stringing on a necklace, and the base bears a brief inscription or cartouche. Some have been used as seals. Pharaohs sometimes commissioned the manufacture of larger images with lengthy inscriptions, such as the commemorative scarab of Queen Tiye. Massive sculptures of scarabs can be seen at Luxor Temple, at the Serapeum in Alexandria (see Serapis) and elsewhere in Egypt.


title: Dung beetle

text: The scarab was of prime significance in the funerary cult of ancient Egypt. Scarabs, generally, though not always, were cut from green stone, and placed on the chest of the deceased. Perhaps the most famous example of such "heart scarabs" is the yellow-green pectoral scarab found among the entombed provisions of Tutankhamen. It was carved from a large piece of Libyan desert glass. The purpose of the "heart scarab" was to ensure that the heart would not bear witness against the deceased at judgement in the Afterlife. Other possibilities are suggested by the "transformation spells" of the "Coffin Texts", which affirm that the soul of the deceased may transform ("xpr") into a human being, a god, or a bird and reappear in the world of the living.


title: Dung beetle

text: In contrast to funerary contexts, some of ancient Egypt's neighbors adopted the scarab motif for seals of varying types. The best-known of these being Judean LMLK seals (8 of 21 designs contained scarab beetles), which were used exclusively to stamp impressions on storage jars during the reign of Hezekiah.


title: Dung beetle

text: The scarab remains an item of popular interest thanks to modern fascination with the art and beliefs of ancient Egypt. Scarab beads in semiprecious stones or glazed ceramics can be purchased at most bead shops, while at Luxor Temple a massive ancient scarab has been roped off to discourage visitors from rubbing the base of the statue "for luck".


title: Dung beetle

text: In Aesop's fable "The Eagle and the Beetle", the eagle kills a hare that has asked for sanctuary with a beetle. The beetle then takes revenge by twice destroying the eagle's eggs. The eagle, in despair, flies up to Olympus and places her latest eggs in Zeus's lap, beseeching the god to protect them. When the beetle finds out what the eagle has done, it stuffs itself with dung, goes straight up to Zeus and flies right into his face. Zeus is startled at the sight of the unpleasant creature, jumping to his feet so that the eggs are broken. Learning of the origin of their feud, Zeus attempts to mediate and, when his efforts to mediate fail, he changes the breeding season of the eagle to a time when the beetles are not above ground.


title: Dung beetle

text: Aristophanes alluded to Aesop's fable several times in his plays. In "Peace", the hero rides up to Olympus to free the goddess Peace from her prison. His steed is an enormous dung beetle which has been fed so much dung that it has grown to monstrous size.


title: Dung beetle

text: Hans Christian Andersen's "The Dung Beetle" tells the story of a dung beetle who lives in the stable of the king's horses in an imaginary kingdom. When he demands golden shoes like those the king's horse wears and is refused, he flies away and has a series of adventures, which are often precipitated by his feeling of superiority to other animals. He finally returns to the stable having decided (against all logic) that it is for him that the king's horse wears golden shoes.


title: Dung beetle

text: In Franz Kafka's "The Metamorphosis", the transformed character of Gregor Samsa is called an "old dung beetle" ("alter Mistkäfer") by a charwoman.


title: Candida albicans

text: Candida albicans is an opportunistic pathogenic yeast that is a common member of the human gut flora. It can also survive outside the human body. It is detected in the gastrointestinal tract and mouth in 40–60% of healthy adults. It is usually a commensal organism, but it can become pathogenic in immunocompromised individuals under a variety of conditions. It is one of the few species of the genus "Candida" that causes the human infection candidiasis, which results from an overgrowth of the fungus. Candidiasis is, for example, often observed in HIV-infected patients.


title: Candida albicans

text: "C. albicans" is the most common fungal species isolated from biofilms either formed on (permanent) implanted medical devices or on human tissue. "C. albicans", "C. tropicalis", "C. parapsilosis", and "C. glabrata" are together responsible for 50–90% of all cases of candidiasis in humans. A mortality rate of 40% has been reported for patients with systemic candidiasis due to "C. albicans". By one estimate, invasive candidiasis contracted in a hospital causes 2,800 to 11,200 deaths yearly in the US. Nevertheless, these numbers may not truly reflect the true extent of damage this organism causes, given new studies indicating that "C. albicans" can cross the blood–brain barrier in mice.


title: Candida albicans

text: "C. albicans" is commonly used as a model organism for fungal pathogens. It is generally referred to as a dimorphic fungus since it grows both as yeast and filamentous cells. However, it has several different morphological phenotypes including opaque, GUT, and pseudohyphal forms. "C. albicans" was for a long time considered an obligate diploid organism without a haploid stage. This is, however, not the case. Next to a haploid stage "C. albicans" can also exist in a tetraploid stage. The latter is formed when diploid "C. albicans" cells mate when they are in the opaque form. The diploid genome size is approximately 29 Mb, and up to 70% of the protein coding genes have not yet been characterized.


title: Candida albicans

text: "C. albicans" is easily cultured in the lab and can be studied both "in vivo" and "in vitro". Depending on the media different studies can be done as the media influences the morphological state of "C. albicans". A special type of medium is CHROMagar "Candida", which can be used to identify different "Candida" species.


title: Candida albicans

text: "Candida albicans" can be seen as a tautology. Candida comes from the Latin word candidus, meaning white. Albicans itself is the present participle of the Latin word albicō, meaning becoming white. This leads to white becoming white, making it a tautology.


title: Candida albicans

text: Over 200 species have been described within the candida genus. The oldest reference to thrush, most likely caused by "C. albicans", dates back to 400 BCE in Hippocrates' work "Of the Epidemics" describing oral candidiasis.


title: Candida albicans

text: The genome of "C. albicans" is almost 16Mb for the haploid size (28Mb for the diploid stage) and consists of 8 sets of chromosome pairs called chr1A, chr2A, chr3A, chr4A, chr5A, chr6A, chr7A and chrRA. The second set ("C. albicans" is diploid) has similar names but with a B at the end. Chr1B, chr2B, ... and chrRB. The whole genome contains 6,198 open reading frames (ORFs). Seventy percent of these ORFs have not yet been characterized. The whole genome has been sequenced making it one of the first fungi to be completely sequenced (next to "Saccharomyces cerevisiae" and "Schizosaccharomyces pombe"). All open reading frames (ORFs) are also available in Gateway-adapted vectors. Next to this ORFeome there is also the availability of a GRACE (gene replacement and conditional expression) library to study essential genes in the genome of "C. albicans". The most commonly used strains to study "C. albicans" are the WO-1 and SC5314 strains. The WO-1 strain is known to switch between white-opaque form with higher frequency while the SC5314 strain is the strain used for gene sequence reference.


title: Candida albicans

text: One of the most important features of the "C. albicans" genome is the high heterozygosity. At the base of this heterozygosity lies the occurrence of numeric and structural chromosomal rearrangements and changes as means of generating genetic diversity by chromosome length polymorphisms (contraction/expansion of repeats), reciprocal translocations, chromosome deletions, Nonsynonymous single-nucleotide polymorphisms and trisomy of individual chromosomes. These karyotypic alterations lead to changes in the phenotype, which is an adaptation strategy of this fungus. These mechanisms are further being explored with the availability of the complete analysis of the "C. albicans" genome.


title: Candida albicans

text: An unusual feature of the genus "Candida" is that in many of its species (including "C. albicans" and "C. tropicalis", but not, for instance, "C. glabrata") the CUG codon, which normally specifies leucine, specifies serine in these species. This is an unusual example of a departure from the standard genetic code, and most such departures are in start codons or, for eukaryotes, mitochondrial genetic codes. This alteration may, in some environments, help these "Candida" species by inducing a permanent stress response, a more generalized form of the heat shock response. However, this different codon usage makes it more difficult to study "C. albicans" protein-protein interactions in the model organism "S. cerevisiae". To overcome this problem a "C. albicans" specific two-hybrid system was developed.


title: Candida albicans

text: The genome of "C. albicans" is highly dynamic, contributed by the different CUG translation, and this variability has been used advantageously for molecular epidemiological studies and population studies in this species. The genome sequence has allowed for identifying the presence of a parasexual cycle (no detected meiotic division) in "C. albicans". This study of the evolution of sexual reproduction in six "Candida" species found recent losses in components of the major meiotic crossover-formation pathway, but retention of a minor pathway. The authors suggested that if "Candida" species undergo meiosis it is with reduced machinery, or different machinery, and indicated that unrecognized meiotic cycles may exist in many species. In another evolutionary study, introduction of partial CUG identity redefinition (from "Candida" species) into "Saccharomyces cerevisiae" clones caused a stress response that negatively affected sexual reproduction. This CUG identity redefinition, occurring in ancestors of "Candida" species, was thought to lock these species into a diploid or polyploid state with possible blockage of sexual reproduction.


title: Candida albicans

text: "C. albicans" exhibits a wide range of morphological phenotypes due to phenotypic switching and bud to hypha transition. The yeast-to-hyphae transition (filamentation) is a rapid process and induced by environmental factors. Phenotypic switching is spontaneous, happens at lower rates and in certain strains up to seven different phenotypes are known. The best studied switching mechanism is the white to opaque switching (an epigenetic process). Other systems have been described as well. Two systems (the high-frequency switching system and white to opaque switching) were discover by David R. Soll and colleagues. Switching in "C. albicans" is often, but not always, influenced by environmental conditions such as the level of CO, anaerobic conditions, medium used and temperature.


title: Candida albicans

text: In its yeast form "C. albicans" ranges from 10 to 12 microns. Spores can form on the pseudohyphae called chlamydospores which survive when put in unfavorable conditions such as dry or hot seasons.


title: Candida albicans

text: Although often referred to as dimorphic, "C. albicans" is, in fact, polyphenic (often also referred to as pleomorphic). When cultured in standard yeast laboratory medium, "C. albicans" grows as ovoid "yeast" cells. However, mild environmental changes in temperature, CO, nutrients and pH can result in a morphological shift to filamentous growth. Filamentous cells share many similarities with yeast cells. Both cell types seem to play a specific, distinctive role in the survival and pathogenicity of "C. albicans". Yeast cells seem to be better suited for the dissemination in the bloodstream while hyphal cells have been proposed as a virulence factor. Hyphal cells are invasive and speculated to be important for tissue penetration, colonization of organs and surviving plus escaping macrophages. The transition from yeast to hyphal cells is termed to be one of the key factors in the virulence of "C. albicans"; however, it is not deemed necessary. When "C. albicans" cells are grown in a medium that mimics the physiological environment of a human host, they grow as filamentous cells (both true hyphae and pseudohyphae). "C. albicans" can also form chlamydospores, the function of which remains unknown, but it is speculated they play a role in surviving harsh environments as they are most often formed under unfavorable conditions.


title: Candida albicans

text: The cAMP-PKA signaling cascade is crucial for the morphogenesis and an important transcriptional regulator for the switch from yeast like cells to filamentous cells is EFG1.


title: Candida albicans

text: Besides the well-studied yeast-to-hyphae transition other switching systems have been described. One such system is the "high-frequency switching" system. During this switching different cellular morphologies (phenotypes) are generated spontaneously. This type of switching does not occur en masse, represents a variability system and it happens independently from environmental conditions. The strain 3153A produces at least seven different colony morphologies. In many strains the different phases convert spontaneously to the other(s) at a low frequency. The switching is reversible, and colony type can be inherited from one generation to another.


title: Candida albicans

text: Being able to switch through so many different (morphological) phenotypes makes "C. albicans" able to grow in different environments, both as a commensal and as a pathogen.


title: Candida albicans

text: In the 3153A strain, a gene called "SIR2" (for silent information regulator), which seems to be important for phenotypic switching, has been found. "SIR2" was originally found in "Saccharomyces cerevisiae" (brewer's yeast), where it is involved in chromosomal silencing—a form of transcriptional regulation, in which regions of the genome are reversibly inactivated by changes in chromatin structure (chromatin is the complex of DNA and proteins that make chromosomes). In yeast, genes involved in the control of mating type are found in these silent regions, and "SIR2" represses their expression by maintaining a silent-competent chromatin structure in this region. The discovery of a "C. albicans SIR2" implicated in phenotypic switching suggests it, too, has silent regions controlled by "SIR2", in which the phenotype-specific genes may reside. How "SIR2" itself is regulated in "S. cerevisiae" may yet provide more clues as to the switching mechanisms of "C. albicans".


title: Candida albicans

text: Next to the dimorphism and the first described high-frequency switching system "C. albicans" undergoes another high-frequency switching process called white to opaque switching, which is another phenotypic switching process in "C. albicans". It was the second high-frequency switching system discovered in "C. albicans". The white to opaque switching is an epigenetic switching system. Phenotypic switching is often used to refer to white-opaque switching, which consists of two phases: one that grows as round cells in smooth, white colonies (referred to as white form) and one that is rod-like and grows as flat, gray colonies (called opaque form). This switch from white cells to opaque cells is important for the virulence and the mating process of "C. albicans" as the opaque form is the mating competent form, being a million times more efficient in mating compared to the white type. This switching between white and opaque form is regulated by the WOR1 regulator (White to Opaque Regulator 1) which is controlled by the mating type locus (MTL) repressor (a1-α2) that inhibits the expression of WOR1. Besides the white and opaque phase there is also a third one: the gray phenotype. This phenotype shows the highest ability to cause cutaneous infections. The white, opaque and gray phenotypes form a tristable phenotypic switching system. Since it is often difficult to differentiate between white, opaque and gray cells phloxine B, a dye, can be added to the medium.


title: Candida albicans

text: A potential regulatory molecule in the white to opaque switching is "Efg1p", a transcription factor found in the WO-1 strain that regulates dimorphism, and more recently has been suggested to help regulate phenotypic switching. "Efg1p" is expressed only in the white and not in the gray cell-type, and overexpression of "Efg1p" in the gray form causes a rapid conversion to the white form.


title: Candida albicans

text: Glucose starvation is a likely common environmental stress encountered by "C. albicans" in its natural habitat. Glucose starvation causes an increase in intracellular reactive oxygen. This stress can lead to mating between two individuals of the same mating type, an interaction that may be frequent in nature under stressful conditions.


title: Candida albicans

text: A very special type of phenotypic switch is the white-GUT switch (Gastrointestinally-IndUced Transition). GUT cells are extremely adapted to survival in the digestive tract by metabolic adaptations to available nutrients in the digestive tract. The GUT cells live as commensal organisms and outcompete other phenotypes. The transition from white to GUT cells is driven by passage through the gut where environmental parameters trigger this transition by increasing the WOR1 expression.


title: Candida albicans

text: "Candida" is found worldwide but most commonly compromises immunocompromised individuals diagnosed with serious diseases such as HIV and cancer. "Candida" are ranked as one of the most common groups of organisms that cause hospital-acquired infections. Especially high-risk individuals are patients that have recently undergone surgery, a transplant or are in the Intensive Care Units (ICU), "C. albicans" infections is the top source of fungal infections in critically ill or otherwise immuncompromised patients. These patients predominantly develop oropharyngeal or thrush candidiasis, which can lead to malnutrition and interfere with the absorption of medication. Methods of transmission include mother to infant through childbirth, people-to-people acquired infections that most commonly occur in hospital settings where immunocompromised patients acquire the yeast from healthcare workers and has a 40% incident rate. People can become infected after having sex with a woman that has an existing vaginal yeast infection. Parts of the body that are commonly infected include the skin, genitals, throat, mouth, and blood. Distinguishing features of vaginal infection include discharge, and dry and red appearance of vaginal mucosa or skin. "Candida" continues to be the fourth most commonly isolated organism in bloodstream infections. Healthy people usually do not suffer (severely) from superficial infections caused by a local alteration in cellular immunity as seen by asthma patients that use oral corticosteroids.


title: Candida albicans

text: It commonly occurs as a superficial infection on mucous membranes in the mouth or vagina. Once in their lives around 75% of women will suffer from vulvovaginal candidiasis (VVC) and about 90% of these infections are caused by "C. albicans". It may also affect a number of other regions. For example, higher prevalence of colonization of "C. albicans" was reported in young individuals with tongue piercing, in comparison to unpierced matched individuals. To infect host tissue, the usual unicellular yeast-like form of "C. albicans" reacts to environmental cues and switches into an invasive, multicellular filamentous form, a phenomenon called dimorphism. In addition, an overgrowth infection is considered a superinfection, the term usually applied when an infection becomes opportunistic and very resistant to antifungals. It then becomes suppressible by antibiotics. The infection is prolonged when the original sensitive strain is replaced by the antibiotic-resistant strain.


title: Candida albicans

text: Candidiasis is known to cause gastrointestinal (GI) symptoms particularly in immunocompromised patients or those receiving steroids (e.g. to treat asthma) or antibiotics. Recently, there is an emerging literature that an overgrowth of fungus in the small intestine of non-immunocompromised subjects may cause unexplained GI symptoms. Small intestinal fungal overgrowth (SIFO) is characterized by the presence of an excessive number of fungal organisms in the small intestine associated with gastrointestinal symptoms. The most common symptoms observed in these patients were belching, bloating, indigestion, nausea, diarrhea, and gas. The underlying mechanism(s) that predisposes to SIFO is unclear. Further studies are needed; both to confirm these observations and to examine the clinical relevance of fungal overgrowth.


title: Candida albicans

text: Systemic fungal infections (fungemias) including those by "C. albicans" have emerged as important causes of morbidity and mortality in immunocompromised patients (e.g., AIDS, cancer chemotherapy, organ or bone marrow transplantation). "C. albicans" often forms biofilms inside the body. Such "C. albicans" biofilms may form on the surface of implantable medical devices or organs. In these biofilms it is often found together with "Staphylococcus aureus". Such multispecies infections lead to higher mortalities. In addition hospital-acquired infections by "C. albicans" have become a cause of major health concerns. Especially once candida cells are introduced in the bloodstream a high mortality, up to 40–60% can occur.


title: Candida albicans

text: Although "Candida albicans" is the most common cause of candidemia, there has been a decrease in the incidence and an increased isolation of non-albicans species of "Candida" in recent years. Preventive measures include maintaining a good oral hygiene, keeping a healthy lifestyle including good nutrition, the careful use of antibiotics, treatment of infected areas and keeping skin dry and clean, free from open wounds.


title: Candida albicans

text: The link between "C. albicans" and Crohn's disease has been investigated in a large cohort. This study demonstrated that members of families with multiple cases of Crohn's disease were more likely to be colonized by "C. albicans" than members of control families. Experimental studies show that chemically-induced colitis promotes "C. albicans" colonization. In turn, "C. albicans" colonization generates anti-"Saccharomyces cerevisiae" antibodies (ASCA), increases inflammation, histological scores and pro-inflammatory cytokine expression.


title: Candida albicans

text: Similarly to antibiotic resistance, resistance to many anti-fungals is becoming a problem. New anti-fungals have to be developed to cope with this problem since only a limited number of anti-fungals are available. A general problem is that in contrast to bacteria, fungi are often overlooked as a potential health problem.


title: Candida albicans

text: Given the fact that candidiasis is the fourth- (to third-) most frequent hospital acquired infection worldwide it leads to immense financial implications. Approximately 60,000 cases of systemic candidiasis each year in the USA alone lead up to a cost to be between $2–4 billion. The total costs for candidiasis are among the highest compared to other fungal infections due to the high prevalence. The immense costs are partly explained by a longer stay in the intensive care unit or hospital in general. An extended stay for up to 21 more days compared to non-infected patients is not uncommon.


title: Candida albicans

text: Gasdermin D (GSDMD) is a protein that in humans is encoded by the GSDMD gene and is a known target of the inflammasome and acts as an effector molecule of programmed cell death known as pyroptosis. This protein determines cell lysis to prevent pathogen replication and results in the release of the inflammatory cytokine interleukin-1β (IL-1β) into the extracellular space to recruit and activate immune cells at the site of infection. Inflammasome activation due to C.albicans infection triggers the release of a cytokine storm necessary to fight the pathogen. Excessive release of these pro-inflammatory mediators has been shown to exaggerate systemic inflammation leading to vascular injury and damage to vital organs. Unfortunately, Candida albicans therapy is often ineffective despite the availability of many antifungal drugs, mainly because of resistance phenomena. During conventional pyroptosis controlled by the inflammasome-GSDMD axis is hijacked by C. albicans to facilitate escape from macrophages through unfolding of hyphae and candidalysin, a fungal toxin released from hyphae. It has been shown that disruption of GSDMD in macrophages infected with Candida albicans reduces the fungal load. In addition, the presence of hyphae and candidalysin are key factors in the activation of GSDMD and the release of Candida from macrophages. Also using Candida-infected mice, inhibition of GSDMD has been shown to paradoxically improve prognosis and survival, indicating that this protein may be a potential therapeutic target in C. albicans-induced sepsis.


title: Candida albicans

text: The biofilm of "C. albicans" is formed in four steps. First, there is the initial adherence step, where the yeast-form cells adhere to the substrate. The second step is called Intermediate step, where the cells propagate to form microcolonies, and germ tubes form to yield hyphae. In the maturation step, the biofilm biomass expands, the extracellular matrix accumulates and drug resistance increases. In the last step of biofilm formation, the yeast-form cells are released to colonize the surrounding environment (dispersion). Yeast cells released from a biofilm have novel properties, including increased virulence and drug tolerance.


title: Candida albicans

text: Zap1, also known as Csr1 and Sur1 (zinc-responsive activator protein), is a transcription factor which is required for the hypha formation in "C. albicans" biofilms. Zap1 controls the equilibrium of yeast and hyphal cells, the zinc transporters and zinc regulated genes in biofilms of "C. albicans".


title: Candida albicans

text: Zinc (Zn) is important for cell function of "C. albicans" and Zap1 controls the Zinc levels in the cells through the zinc transporters Zrt1 and Zrt2. The regulation of zinc concentration in the cells is important for the cell viability and if the zinc levels get too high, it is toxic for the cells. The Zrt1 is transporting the zinc ions with high affinity and the Zrt2 is transporting the zinc ions with low affinity.


title: Candida albicans

text: The ability to switch between yeast cells and hyphal cells is an important virulence factor. Many proteins play a role in this process.


title: Candida albicans

text: Filamentation in "C. albicans" is a very complex process. The formation of hyphae can for example help "Candida albicans" to escape from macrophages in the human body. Moreover, "C. albicans" undergo yeast-to-hyphal transition within the acidic macrophage phagosome. This initially causes phagosome membrane distension which eventually leads to phagosomal alkalinization by physical rupture, followed by escape.


title: Candida albicans

text: Hwp1 stands for Hyphal wall protein 1. Hwp1 is a mannoprotein located on the surface of the hyphae in the hyphal form of "C. albicans". Hwp1 is a mammalian transglutaminase substrate. This host enzyme allows "Candida albicans" to attach stably to host epithelial cells. Adhesion of "C. albicans" to host cells is an essential first step in the infection process for colonization and subsequent induction of mucosal infection.


title: Candida albicans

text: The RNA-binding protein Slr1 plays a role in instigating hyphal formation and virulence in "C. albicans".


title: Candida albicans

text: Candidalysin is a cytolytic 31-amino acid α-helical peptide toxin that is released by "C. albicans" during hyphal formation. It contributes to virulence during mucosal infections.


title: Candida albicans

text: Due to its nature as a model organism, being an important human pathogen and the alternative codon usage (CUG translated into serine rather than leucine), several specific projects and tools have been created to study "C. albicans". The diploid nature and the absence of a sexual cycle, however, makes it a hard to study organism. In the last 20 years, however, many systems have been developed to study "C. albicans" in a more in depth genetic level.


title: Candida albicans

text: The most used selection markers in "C. albicans" are the CaNAT1 resistance marker (confers resistance against nourseothricin) and MPAr or IMH3r (confers resistance to mycophenolic acid).


title: Candida albicans

text: Next to the above-mentioned selection makers a few auxotrophic strains were generated to work with auxotrophic makers. The URA3 marker (URA3 blaster method) is an often-used strategy in uridine auxotrophic strains; however, studies have shown that differences in URA3 position in the genome can be involved in the pathogeny of "C. albicans". Besides the URA3 selection one can also use the histidine, leucine and arginine autotrophy. The advantage of using those autotrophies lies in the fact that they exhibit wild-type or nearly wild-type virulence in a mouse model compared to the URA3 system. One application of the leucine, arginine and histidine autotrophy is for example the candida two-hybrid system.


title: Candida albicans

text: The full genome of "C. albicans" has been sequenced and made publicly available in a Candida database. The heterozygous diploid strain used for this full genome sequence project is the laboratory strain SC5314. The sequencing was done using a whole-genome shotgun approach.


title: Candida albicans

text: Every predicted ORF has been created in a gateway adapted vector (pDONR207) and made publicly available. The vectors (plasmids) can be propagated in "E.coli" and grown on LB+gentamicin medium. This way every ORF is readily available in an easy to use vector. Using the gateway system it is possible to transfer the ORF of interest to any other gateway adapted vector for further studies of the specific ORF.


title: Candida albicans

text: Contrary to the yeast "S. cerevisiae" episomal plasmids do not stay stable in "C. albicans". In order to work with plasmids in "C. albicans" an integrative approach (plasmid integration into the genome) thus has to be used. A second problem is that most plasmid transformations are rather inefficient in "C. albicans"; however, the CIp10 plasmid overcomes these problems and can be used with ease to transform "C. albicans" in a very efficient way. The plasmid integrates inside the RP10 locus as disruption of one RP10 allele does not seem to affect the viability and growth of "C. albicans". Several adaptations of this plasmid have been made after the original became available.


title: Candida albicans

text: Due to the aberrant codon usage of "C. albicans" it is less feasible to use the common host organism ("Saccharomyces cerevisiae") for two-hybrid studies. To overcome this problem a "C. albicans" two-hybrid (C2H) system was created. The strain SN152 that is auxotrophic for leucine, arginine and histidine was used to create this C2H system. It was adapted by integrating a HIS1 reporter gene preceded by five LexAOp sequences.


title: Candida albicans

text: In the C2H system the bait plasmid (pC2HB) contains the "Staphylococcus aureus" LexA BD, while the prey plasmid (pC2HP) harbors the viral AD VP16. Both plasmids are integrative plasmids since episomal plasmids do not stay stable in "C. albicans". The reporter gene used in the system is the "HIS1" gene. When proteins interact, the cells will be able to grow on medium lacking histidine due to the activation of the "HIS1" reporter gene. Several interactions have thus far been detected using this system in a low scale set up. A first high-throughput screening has also been performed. Interacting proteins can be found at the BioGRID.


title: Candida albicans

text: Besides the C2H system, a BiFC system has been developed to study protein-protein interactions in "C. albicans". With this systems protein interactions can be studied in their native sub cellular location contrary to a C2H system in which the proteins are forced into the nucleus. With BiFC one can study for example protein interactions that take place at the cell membrane or vacuolar membrane.


title: Candida albicans

text: Both DNA and protein microarrays were designed to study DNA expression profiles and antibody production in patients against "C. albicans" cell wall proteins.


title: Candida albicans

text: Using a tetracycline-regulatable promoter system a gene replacement and conditional expression (GRACE) library was created for 1,152 genes. By using the regulatable promoter and having deleted 1 of the alleles of the specific gene it was possible to discriminate between non-essential and essential genes. Of the tested 1,152 genes 567 showed to be essential. The knowledge on essential genes can be used to discover novel antifungals.


title: Candida albicans

text: CRISPR/Cas9 has been adapted to be used in "C. albicans". Several studies have been performed using this system.


title: Candida albicans

text: "C. albicans" has been used in combination with carbon nanotubes (CNT) to produce stable electrically conductive bio-nano-composite tissue materials that have been used as temperature-sensing elements.


title: Martin Eberhard

text: Martin Eberhard (born ) is an American inventor, engineer and entrepreneur who co-founded Tesla, Inc. (then Tesla Motors) with Marc Tarpenning in 2003. Eberhard served as Tesla's original chairman, and its CEO until late 2007. In 2015, he was inducted into the University of Illinois Engineering Hall of Fame.


title: Martin Eberhard

text: Eberhard grew up in Kensington, California, a community in the Berkeley Hills. He received a B.S. in computer engineering from the University of Illinois Urbana-Champaign in 1982 and an M.S. in electrical engineering from the same school in 1984.


title: Martin Eberhard

text: Eberhard began his career as an electrical engineer at Wyse Technology, where he designed the WY-30 ASCII computer terminal as his first product. Eberhard co-founded Network Computing Devices in 1987, where he served as chief engineer through its IPO on 1992.


title: Martin Eberhard

text: In 1996, Eberhard founded NuvoMedia with Marc Tarpenning, where they developed the Rocket eBook, the first e-book with secure internet delivery of content. Eberhard served as chairman and CEO until NuvoMedia’s acquisition by Gemstar in 2000.


title: Martin Eberhard

text: Eberhard's interest in sports cars, concern about the dependence on oil imports, and global warming led him to think about EV cars as part of the solution. He found out about and attempted to purchase a prototype EV sports car called the AC Propulsion tzero. He provided some financial and technical support in the conversion of the tzero to lithium-ion batteries. Eberhard then tried to convince AC Propulsion founder Alan Cocconi to turn the tzero into a production car. When Cocconi declined, Eberhard co-founded (with Marc Tarpenning), and became the first CEO of, Tesla Motors, an electric car company in Menlo Park, California in 2003.


title: Martin Eberhard

text: 1) An electric car should not be a compromise. With the right technology choices, it is possible to build electric cars that are actually better cars than their competition.


title: Martin Eberhard

text: 2) Battery technology is key to a successful electric car. Lithium ion batteries are not only suitable of automotive use; they are game-changing, making decent driving range a reality.


title: Martin Eberhard

text: 3) If designed right, electric cars can appeal to even the most serious car enthusiast, as electric drive is capable of seriously outperforming internal combustion engines.


title: Martin Eberhard

text: Eberhard drives the second of Tesla Motors Founder's Series Roadsters cars, which is the first series of the Tesla Roadster (2008). The Tesla Roadster is a battery electric sportscar with (EPA) range.


title: Martin Eberhard

text: On November 30, 2007, Tesla released a press release titled "Martin Eberhard, Co-founder of Tesla Motors, to Transition to Advisory Board." Fortune magazine reported in December 2007 that chairman Elon Musk had asked Eberhard to leave. Musk stated in an interview that it was not due to ideological differences, but that he did not see a role for Eberhard. On January 7, 2008, the New York Times reported that Tesla Motors issued a statement explaining that the co-founder and former chief executive, Martin Eberhard, "has transitioned from the board of directors and executive management of the company to the advisory board."


title: Martin Eberhard

text: Eberhard noted that while he had signed a non-disclosure agreement with Tesla, "so I must, by contract, be a bit careful about how I word things", he was not happy with the transition. In his since-deleted Tesla Founders Blog, Eberhard criticized Tesla layoffs, which he labeled a "stealth bloodbath".


title: Martin Eberhard

text: In June 2009, Eberhard brought a lawsuit against Elon Musk for libel, slander, and breach of contract, alleging that Musk pushed him out of the company, publicly disparaged him, and compromised Tesla's financial health. In August 2009, Eberhard dropped the lawsuit for undisclosed reasons. A Tesla spokesperson declined to comment on the change, raising the likelihood of a settlement. In September 2009, Tesla confirmed the settlement, but did not provide further details.


title: Martin Eberhard

text: Eberhard confirmed in an interview with CNBC in October 2019 that he is still a shareholder of Tesla, and is still rooting for their success.


title: Martin Eberhard

text: In 2010, Eberhard confirmed to Autoblog Green that he was doing work with Volkswagen, but no further details were provided.


title: Martin Eberhard

text: In September 2016, Eberhard founded stealth-mode startup inEVit in a bid to supply major OEMs with electric drivetrains and power storage solutions.


title: Martin Eberhard

text: SF Motors (now Seres) acquired inEVit in October 2017. Eberhard served as chief innovation officer until leaving in July 2018.


title: Springfield, Massachusetts

text: Springfield is a city in the Commonwealth of Massachusetts, United States, and the seat of Hampden County. Springfield sits on the eastern bank of the Connecticut River near its confluence with three rivers: the western Westfield River, the eastern Chicopee River, and the eastern Mill River. At the 2020 census, the city's population was 155,929, making it the third-largest city in Massachusetts, the fourth-most populous city in New England after Boston, Worcester, and Providence, and the 12th-most populous in the Northeastern United States. Metropolitan Springfield, as one of two metropolitan areas in Massachusetts (the other being Greater Boston), had a population of 699,162 in 2020.


title: Springfield, Massachusetts

text: Springfield was founded in 1636, the first Springfield in the New World. In the late 1700s, during the American Revolution, Springfield was designated by George Washington as the site of the Springfield Armory because of its central location. Subsequently it was the site of Shays' Rebellion. The city would also play a pivotal role in the Civil War, as a stop on the Underground Railroad and home of abolitionist John Brown, widely known for his raid on Harpers Ferry, and for the Armory's manufacture of the famed "Springfield rifles" used ubiquitously by Union troops. Closing during the Johnson administration, today the national park site features the largest collection of historic American firearms in the world.


title: Springfield, Massachusetts

text: Today the city is the largest in western New England, and the urban, economic, and media capital of Massachusetts' section of the Connecticut River Valley, colloquially known as the Pioneer Valley.


title: Springfield, Massachusetts

text: Springfield has several nicknames—"The City of Firsts", due to the many innovations developed there, such as the first American dictionary, the first American gas-powered automobile, and the first machining lathe for interchangeable parts; "The City of Homes", due to its Victorian residential architecture; and "Hoop City", as basketball was invented in Springfield in 1891 by Canadian James Naismith.Also, the city gained a new nickname when not one night could go by without the sound of gunshots or police sirens being heard. Their new nickname is "Los Angeles 2".


title: Springfield, Massachusetts

text: Hartford, the capital of Connecticut, lies south of Springfield, on the western bank of the Connecticut River. The Hartford–Springfield region is known as the Knowledge Corridor because it hosts over 160,000 university students and over 32 universities and liberal arts colleges—the second-highest concentration of higher-learning institutions in the United States. The city of Springfield itself is home to Springfield College, Western New England University, American International College, and Springfield Technical Community College, among other higher educational institutions.


title: Springfield, Massachusetts

text: Springfield was founded in 1636 by English Puritan William Pynchon as "Agawam Plantation" under the administration of the Connecticut Colony. In 1641 it was renamed after Pynchon's hometown of Springfield, Essex, England, following incidents, including trade disputes as well as Captain John Mason's hostilities toward native tribes, which precipitated the settlement's joining the Massachusetts Bay Colony. During its early existence, Springfield flourished both as an agricultural settlement and as a trading post, although its prosperity waned dramatically during (and after) King Philip's War in 1675, when natives laid siege to it and burned it to the ground as part of the ongoing campaign. During that attack, three-quarters of the original settlement was burned to the ground, with many of Springfield's residents survived by taking refuge in John Pynchon's brick house, the "Old Fort", the first such house to be built in the Connecticut River Valley. Out of the siege, Miles Morgan and his sons were lauded as heroes; as one of the few homesteads to survive the attack, alerting troops in Hadley, as well as Toto, often referred to as the "Windsor Indian" who, running 20 miles from Windsor, Connecticut to the settlement, was able to give advance warning of the attack.


title: Springfield, Massachusetts

text: The original settlement—today's downtown Springfield—was located atop bluffs at the confluence of four rivers, at the nexus of trade routes to Boston, Albany, New York City, and Montreal, and with some of the northeastern United States' most fertile soil. In 1777, Springfield's location at numerous crossroads led George Washington and Henry Knox to establish the United States' National Armory at Springfield, which produced the first American musket in 1794, and later the famous Springfield rifle. From 1777 until its closing during the Vietnam War, the Springfield Armory attracted skilled laborers to Springfield, making it the United States' longtime center for precision manufacturing. The near-capture of the armory during Shays' Rebellion of 1787 led directly to the formation of the U.S. Constitutional Convention.


title: Springfield, Massachusetts

text: During the 19th and 20th centuries, Springfielders produced many innovations, including the first American-English dictionary (1805, Merriam-Webster); the first use of interchangeable parts and the assembly line in manufacturing (1819, Thomas Blanchard); the first American horseless car (1825, Thomas Blanchard); the mass production of vulcanized rubber (1844, Charles Goodyear); the first American gasoline-powered car (1893, Duryea Brothers); the first successful motorcycle company (1901, "Indian"); one of America's first commercial radio stations (1921, WBZ, broadcast from the Hotel Kimball); and most famously, the world's second-most-popular sport, basketball (1891, Dr. James Naismith). Springfield would play major roles in machine production, initially driven by the arms industry of the Armory, as well as from private companies such as Smith & Wesson, established by Horace Smith and Daniel B. Wesson. Similarly, the industrial economy led Thomas and Charles Wason to establish the Wason Manufacturing Company, which produced the first manufactured sleeping car. The largest railcar works in New England, Wason produced 100 cars a day at its peak; the company was eventually was purchased by Brill in 1907 and closed during the Depression in 1937. Among numerous other industries, during the first half of the 20th century Springfield also produced brass goods, chemicals, clothing and knit goods, paper goods, watches, boilers, engines, manufacturing machinery, silverware, jewelry, skates, carriages, buttons, needles, toys, and printed books and magazines.


title: Springfield, Massachusetts

text: Springfield underwent a protracted decline during the second half of the 20th century, due largely to the decommissioning of the Springfield Armory in 1969; poor city planning decisions, such as the location of the elevated I-91 along the city's Connecticut River front; and overall decline of industry throughout the northeastern United States. During the 1980s and 1990s, Springfield developed a national reputation for crime, political corruption, and cronyism. During the early 21st century, Springfield saw long-term revitalization projects and several large projects, including the $1 billion New Haven–Hartford–Springfield intercity rail; a $1 billion MGM casino.


title: Springfield, Massachusetts

text: According to the United States Census Bureau, the city has a total area of , of which are land and , or 3.65%, are water. Once nicknamed "The City in a Forest", Springfield features over of urban parkland, 12% of its total land area.


title: Springfield, Massachusetts

text: Located in the fertile Connecticut River Valley, surrounded by mountains, bluffs, and rolling hills in all cardinal directions, Springfield sits on the eastern bank of the Connecticut River, near its confluence with two major tributary rivers—the western Westfield River, which flows into the Connecticut opposite Springfield's South End Bridge; and the eastern Chicopee River, which flows into the Connecticut less than north of Springfield, in the city of Chicopee (which constituted one of Springfield's most populous neighborhoods until it separated and became an independent municipality in 1852). The Connecticut state line is only south of Springfield, beside the wealthy suburb of Longmeadow, which itself separated from Springfield in 1783.


title: Springfield, Massachusetts

text: Springfield's densely urban Metro Center district surrounding Main Street is relatively flat, and follows the north–south trajectory of the Connecticut River; however, as one moves eastward, the city becomes increasingly hilly.


title: Springfield, Massachusetts

text: Aside from its rivers, Springfield's second most prominent topographical feature is the city's Forest Park. Forest Park lies in the southwestern corner of the city, surrounded by Springfield's attractive garden districts, Forest Park and Forest Park Heights, which feature over 600 Victorian Painted Lady mansions. Forest Park also borders Western Massachusetts' most affluent town, Longmeadow. Springfield shares borders with other well-heeled suburbs such as East Longmeadow, Wilbraham, Ludlow and the de-industrializing city of Chicopee. The small cities of Agawam and West Springfield lie less than a mile (1.6 km) from Springfield's Metro Center, across the Connecticut River.


title: Springfield, Massachusetts

text: The City of Springfield also owns the Springfield Country Club, located in the autonomous city of West Springfield, which separated from Springfield in 1774.


title: Springfield, Massachusetts

text: Springfield, like other cities in southern New England, has a hot-summer humid continental climate (Köppen: "Dfa") with four distinct seasons and precipitation evenly distributed throughout the year, but the intensity (and sometimes the duration) of warmer periods greater than northern areas. Winters are cold with a daily average in January of around . During winter, nor'easter storms can drop significant snowfalls on Springfield and the Connecticut River Valley. Temperatures below can occur each year, though the area does not experience the high snowfall amounts and blustery wind averages of nearby cities such as Worcester, Massachusetts and Albany, New York.


title: Springfield, Massachusetts

text: Springfield's summers are very warm and sometimes humid. During summer, several times per month, on hot days afternoon thunderstorms will develop when unstable warm air collides with approaching cold fronts. The daily average in July is around . Usually several days during the summer exceed , constituting a "heat wave". Spring and fall temperatures are usually pleasant, with mild days and crisp, cool nights. Precipitation averages annually, and snowfall averages , most of which falls from mid-December to early March. Although not unheard of, extreme weather events like hurricanes and tornadoes occur infrequently in Springfield compared with other areas in the country. On the occasions that hurricanes have hit New England, Springfield's inland, upriver location has caused its damages to be considerably less than shoreline cities like New Haven, Connecticut and Providence, Rhode Island.


title: Springfield, Massachusetts

text: On June 1, 2011, Springfield was directly struck by the second-largest tornado ever to hit Massachusetts. With wind speeds exceeding , the tornado left three dead, hundreds injured, and over 500 homeless in the city alone. The tornado caused hundreds of millions of dollars worth of damage to Springfield and destroyed nearly everything in a path from Westfield to Charlton, Massachusetts. It was the first deadly tornado to strike Massachusetts since May 29, 1995.


title: Springfield, Massachusetts

text: According to the 2010 Census, Springfield had a population of 153,060, of which 72,573 (47.4%) were male and 80,487 (52.6%) were female. 73.0% of the population were over 18 years old, and 10.9% were over 65 years old; the median age was 32.2 years. The median age for males was 30.2 years and 34.1 years for females.


title: Springfield, Massachusetts

text: According to the 2010 Census, there were 61,706 housing units in Springfield, of which 56,752 were occupied. This was the highest average of home occupancy among the four distinct Western New England metropolises (the other three being Hartford, New Haven, and Bridgeport, Connecticut). Also as of 2010, Springfield features the highest average homeowner occupancy ratio among the four Western New England metropolises at 50%—73,232 Springfielders live in owner-occupied units, versus 74,111 in rental units. By comparison, as of the 2010 Census, New Haven features an owner occupancy rate of 31%; Hartford of 26%; and Bridgeport of 43%.


title: Springfield, Massachusetts

text: In terms of race and ethnicity, Springfield is 51.8% White, 22.3% Black or African American, 0.6% American Indian and Alaska Native, 2.4% Asian (1.2% Vietnamese, 0.3% Chinese, 0.2% Indian, 0.1% Cambodian, 0.1% Filipino, 0.1% Korean, 0.1% Pakistani, 0.1% Laotian), 0.1% Native Hawaiian and Other Pacific Islander, 18.0% from Some Other Race, and 4.7% from Two or More Races (1.5% White and Black or African American; 1.0% White and Some Other Race). Hispanics and Latinos of any race made up 38.8% of the population (33.2% Puerto Rican, 1.7% Dominican, 1.0% Mexican, 0.5% Guatemalan, 0.3% Cuban, 0.2% Colombian, 0.2% Spanish, 0.2% Salvadoran, 0.1% Peruvian, 0.1% Ecuadorian, 0.1% Panamanian, 0.1% Costa Rican, 0.1% Honduran). Non-Hispanic Whites were 36.7% of the population in 2010, down from 84.1% in 1970.


title: Springfield, Massachusetts

text: In 2010, Springfield ranked 35th in the United States' City Crime Rankings—its second-lowest ranking in recent years, (in 2009, it ranked 51st). Springfield's 2010 crime rating of 142 is down approximately 50% from its heights in the late 1990s and 2000s.


title: Springfield, Massachusetts

text: The Urban Land Institute stated in 2010 that "the perception of crime [in Springfield] appears to be worse than the reality".


title: Springfield, Massachusetts

text: By another measure, crime and population data collected by the FBI, and indexed by NeighborhoodScout showed between 2010 and 2018 the violent crime rate for Springfield decreased by approximately 52.5%, whereas the property crime rate declined by 54%; both rates remain more than twice their respective state averages, as of 2018.


title: Springfield, Massachusetts

text: Springfield's vicinity to both Boston and New York City lends it a location well suited for distribution, and in the past this has played a significant role in its economy. For this reason in the early 20th century it was the largest producer in New England of commercially produced cakes and pastries, and among the largest in bread—with one 1926 estimate of 1.4 million loaves of bread and 14 million breakfast rolls produced in the city on a weekly basis.


title: Springfield, Massachusetts

text: Today Springfield's top five industries (in order, by number of workers) are: Education and Health Services; Trade and Transportation; Manufacturing; Tourism and Hospitality; and Professional & Business Services. Springfield is considered to have a "mature economy", which protects the city to a degree during recessions and inhibits it somewhat during bubbles. Springfield is considered to have one of America's top emerging multi-cultural markets—the city features a 33% Latino population with buying power that has increased over 295% from 1990 to 2006. As of 2006, more than 60% of Hispanic Springfielders had arrived in the city since 1986.


title: Springfield, Massachusetts

text: With 25 universities and colleges within a radius from Springfield, including several of America's most prestigious universities and liberal arts colleges, and more than six institutions within the city itself, the Hartford–Springfield metropolitan area has been dubbed the Knowledge Corridor by regional educators, civic authorities, and businessmen—touting its 32 universities and liberal arts colleges, numerous highly regarded hospitals, and nearly 120,000 students. The Knowledge Corridor universities and colleges provide the region with an educated workforce, which yields a yearly GDP of over $100 billion—more than at least 16 U.S. States. Hartford–Springfield has become home to a number of biotech firms and high-speed computing centers. As of 2009 Springfield ranks as the 24th most important high-tech center in the United States with approximately 14,000 high-tech jobs.


title: Springfield, Massachusetts

text: In 2010, the median household income was $35,236. Median income for the family was $51,110. The per capita income was $16,863. About 21.3% of families and 26.8% of the population were below the poverty line, including 40.0% of those under age 18 and 17.5% of those age 65 or over.


title: Springfield, Massachusetts

text: The City of Springfield is the economic center of Western Massachusetts. It features the Pioneer Valley's largest concentration of retail, manufacturing, entertainment, banking, legal, and medical groups. Springfield is home to the Commonwealth of Massachusetts' largest Fortune 100 company, MassMutual Financial Group. It is also home to the world's largest producer of handguns, Smith & Wesson, founded in 1852. It is home to Merriam Webster, the first and most widely read American–English dictionary, founded in 1806. It also serves as the headquarters of the professional American Hockey League, the NHL's minor league, Peter Pan Bus, and Big Y Supermarkets, among other businesses.


title: Springfield, Massachusetts

text: Springfield is also home to the Commonwealth of Massachusetts' third largest employer, Baystate Health, with over 10,000 employees. Baystate is the western campus of Tufts University School of Medicine. Baystate Health is in the midst of a $300 million addition; nicknamed "The Hospital of the Future", it is the largest construction project in New England. In addition to Baystate, Springfield features two other nationally ranked hospitals; Mercy Medical, run by The Sisters of Providence, and Shriners Hospital for Children. The following companies maintain their headquarters in Springfield:


title: Springfield, Massachusetts

text: With a history spanning nearly four centuries, Springfield has been home to a number of legacy companies that were household names, including:


title: Springfield, Massachusetts

text: Springfield's mosaic of ethnic communities have long played a role in its culinary institutions, with many newer smaller restaurants and several decades-old establishments in its downtown. Among its oldest institutions are Smith's Billiards, founded in 1902, which serves Theodore's Blues, Booze, & BBQ, a neighboring barbecue and music venue since 1979, recognized as the "Best Blues Club in the Country" in 2004 by the Blues Foundation. Other venues include The Student Prince Cafe, a long-running German restaurant at its downtown location since 1935, featured on "Thrillist's" 2014 Editor's Choice bars, and a wide array of long-running acclaimed Italian restaurants such as Frigo's (1950), The Red Rose (1963), and Leone's (1988) and pastry shops like La Fiorentina (1947).


title: Springfield, Massachusetts

text: In recent decades the city has also become home to a number of cuisines not found elsewhere in the region, including Cajun restaurant Chef Wayne's Big Mamou, Lebanese restaurant Nadim's Downtown, local chain Puerto Rican Bakery, and a host of other Greek, Jamaican, Mexican, and Vietnamese venues.


title: Springfield, Massachusetts

text: Among the best known of Springfield's eateries outside the city is Friendly's, the East Coast restaurant chain whose first store was opened by the Blake brothers as "Friendly Ice Cream" in Springfield in 1935; their original location has since closed and today the chain maintains its headquarters in the neighboring town of Wilbraham.


title: Springfield, Massachusetts

text: Springfield is home to five distinct museums at the Quadrangle, along with the ornate Springfield City Library—an architecturally significant example of the City Beautiful movement. The Quadrangle's five distinct collections include the first American-made planetarium, designed and built (1937) by Frank Korkosz; the Dr Seuss National Memorial Sculpture Garden; the largest collection of Chinese cloisonne outside of China; and the original casting of Augustus Saint Gaudens's most famous sculpture, "Puritan".


title: Springfield, Massachusetts

text: The Quadrangle's five museums are the Museum of Fine Arts, which features a large Impressionist collection; the George Walter Vincent Smith Art Museum, a collection of Asian curiosities; the Springfield Science Museum, which features a life-size Tyrannosaurus Rex, an aquarium and the United States' first planetarium; the Amazing World of Dr. Seuss Museum, as Springfield is the birthplace of Theodor Geisel; and the Lyman and Merrie Wood Museum of Springfield History, a museum about the multi-faceted city.


title: Springfield, Massachusetts

text: In 2017 the Amazing World of Dr. Seuss Museum opened in the former location, the William Pynchon Memorial Building. The collections of the Connecticut Valley Historical Museum are now located in the Lyman and Merrie Wood Museum of Springfield History.


title: Springfield, Massachusetts

text: Springfield's Indian Orchard neighborhood is home to the RMS Titanic Historical Society's Titanic Museum, which displays a collection of rare artifacts that tell stories about the ill-fated ocean liner's passengers and crew.


title: Springfield, Massachusetts

text: In 2011, Springfield's music scene was eclectic. It featured a notable heavy rock scene, from which the bands Gaiah, Staind, All That Remains, Shadows Fall, and The Acacia Strain rose to national prominence. Jazz and blues rival rock in popularity.


title: Springfield, Massachusetts

text: In 2010 and 2011 the Springfield-headquartered Hampden Bank sponsored the Hoops City Jazz & Art Festival, a three-day event that drew approximately 30,000 people to Metro Center to hear varieties of different jazz music—from smooth jazz, to hard bop, to New Orleans-style jazz. Headliners included Springfield great Taj Mahal, the Average White Band, and Poncho Sanchez.


title: Springfield, Massachusetts

text: In addition to its nickname "The City of Firsts", Springfield is known as "The City of Homes" for its attractive architecture, which differentiates it from most medium-size, Northeastern American cities. Most of Springfield's housing stock consists of Victorian "Painted Ladies" (similar to those found in San Francisco); however, Springfield also features Gilded Age mansions, urban condominiums buildings, brick apartment blocks, and more suburban post-World War II architecture (in the Sixteen Acres and Pine Point neighborhoods). While Springfield's architecture is attractive, much of its built-environment stems from the 19th and early 20th centuries when the city experienced a period of "intense and concentrated prosperity"—today, its Victorian architecture can be found in various states of rehabilitation and disrepair. As of 2011, Springfield's housing prices are considerably lower than nearby New England cities which do not feature such intricate architecture.


title: Springfield, Massachusetts

text: In Metro Center, stands the two prominent skyscrapers, Tower Square, and Monarch Place, the latter of which is the tallest building in Massachusetts outside of Boston. Many of the city's Victorian buildings including former hotels, factories, and other institutions have been converted into apartment buildings and luxury condominiums. For example, Springfield's ornate Classical High School (235 State Street), with its immense Victorian atrium—where Dr. Seuss, Timothy Leary, and Taj Mahal all went to high school—is now a luxury condominium building. The Hotel Kimball, (140 Chestnut Street), which hosted several U.S. Presidents as guests and once featured the United States' first commercial radio station (WBZ), has been converted into The Kimball Towers Condominiums. The former McIntosh Shoe Company (158 Chestnut Street), one of Springfield's finest examples of the Chicago School of Architecture, has been converted into industrial-style condominiums; and the red-brick, former Milton Bradley toy factory is now Stockbridge Court Apartments (45 Willow Street). In the Ridgewood Historic District, the 1950s-futurist Mulberry House (101 Mulberry Street), is now a condominium building that features some of the finest views of Springfield.


title: Springfield, Massachusetts

text: Forest Park (and Forest Park Heights), surrounding by the Forest Park, is a New England Garden District featuring more than 600 Victorian Painted Ladies. The McKnight National Historic District, America's first planned residential neighborhood, (1881), features more than 900 Victorian Painted Ladies, many of which have been rehabilitated by Springfield's growing LGBT community. The Old Hill, Upper Hill, and Bay neighborhoods also feature this type of architecture.


title: Springfield, Massachusetts

text: Maple High, which is architecturally (and geographically) distinct from, but often included with Springfield's economically depressed Six Corners neighborhood, was Springfield's first "Gold Coast". Many mansions from the early 19th century and later gilded age stand atop a bluff on Maple Street, overlooking the Connecticut River. The Ridgewood Historic district on Ridgewood and Mulberry Streets also feature historic mansions from the 19th and early 20th centuries.


title: Springfield, Massachusetts

text: Springfield—like many mid-size Northeastern cities, e.g., Hartford, Albany, and New Haven—from the 1950s to the 1970s, razed a significant number of historic commercial buildings in the name of urban renewal. In 1961, this included Unity Church, the first building designed by the young Henry Hobson Richardson. Springfield's Metro Center remains more aesthetically cohesive than many its peer cities; however, as elsewhere, the city currently features a patchwork of parking lots and grand old buildings. Current efforts are underway to improve the cohesion of Springfield's Metro Center, including the completed Main Street and State Street Corridor improvement projects, the upcoming $70 million renovation to Springfield's 1926 Union Station and the renovation of the Epiphany Tower on State Street into a new hotel. New constructions include the architecturally award-winning, $57 million Moshe Safdie-designed Federal Building on State Street.


title: Springfield, Massachusetts

text: Efforts to establish a public library began in the 1850s. In fiscal year 2008, the city of Springfield spent 1.13% ($5,321,151) of its budget on its public library—approximately $35 per person, per year ($46.12 adjusted for inflation to 2022). In fiscal year 2009, Springfield spent about 1% ($5,077,158) of its budget on the library—approximately $32 per person, per year ($42.32 adjusted for inflation to 2022). Springfield has Massachusetts' 2nd largest library circulation, behind Boston.


title: Springfield, Massachusetts

text: As of 2012, the public library purchases access for its patrons to databases owned by the following companies:


title: Springfield, Massachusetts

text: Besides Springfield's historic connection with basketball, the city has a rich sporting history. Volleyball was invented in the adjacent city of Holyoke, and the first exhibition match was held in 1896 at the International YMCA Training School, now known as Springfield College.


title: Springfield, Massachusetts

text: Ice hockey has been played professionally in Springfield since the 1920s, and Springfield is home to the league headquarters of the American Hockey League. The Springfield Indians of the American Hockey League (now located in Utica, New York) was the oldest minor league hockey franchise in existence. In 1994 the team relocated to Worcester and was replaced by the Springfield Falcons, who played at the MassMutual Center. The Falcons were then replaced by the Springfield Thunderbirds in 2016. For parts of two seasons (1978–1980) the NHL Hartford Whalers played in Springfield while their arena was undergoing repairs after a roof collapse. On the amateur level, the Junior A Springfield Olympics played for many years at the Olympia, while American International College's Yellow Jackets compete in NCAA Division I hockey.


title: Springfield, Massachusetts

text: Basketball remains a popular sport in Springfield's sporting landscape. Springfield is currently home to the Western Mass Zombies a minor league basketball team participating in the East Coast Basketball League. The Zombies in 2021 were named ECBL Mid-Atlantic Champions. Prior to the 2014–2015 season, Springfield was home to the Springfield Armor of the NBA Development League, which began play in 2009 at the MassMutual Center. Beginning in the 2011–2012 season, the Armor was the exclusive affiliate of the Brooklyn Nets. For many years, the Hall of Fame Tip-Off Classic has been the semi-official start to the college basketball season, and the NCAA Division II championships are usually held in Springfield. The Metro Atlantic Athletic Conference will play its championships in Springfield from 2012 to 2014. The New England Blizzard of the ABL played its first game in Springfield, and several minor pro men's and women's teams have called the city home, including the Springfield Fame of the United States Basketball League (the league's inaugural champion in 1985) and the Springfield Hall of Famers of the Eastern Professional Basketball League.


title: Springfield, Massachusetts

text: Springfield has had professional baseball in the past, and according to its current mayor, remains intent on pursuing it in the future. The Springfield Giants of the Single- and Double-A Eastern League played between 1957 and 1965. The team was quite successful, winning consecutive championships in 1959, 1960 and 1961, by startling coincidence the same seasons in which the Springfield Indians won three straight Calder Cup championships in hockey. The Giants played at Pynchon Park by the Connecticut River until relocating after the 1965 season. Pynchon Park's grandstands were destroyed by fire the year after in 1966. Before that time, the Springfield Cubs played in the minor league New England League from 1946 until 1949, after which the league folded; they then played in the International League until 1953. For many years before the Giants, Springfield was also a member of the Eastern League, between 1893 and 1943. In general, the team was named the Ponies, but it also carried the nicknames of "Maroons" (1895), "Green Sox" (1917), "Hampdens" (1920–1921), "Rifles (1932, 1942–1943) and "Nationals" (1939–1941). The team located closest are the Valley Blue Sox of the New England Collegiate Baseball League who play their games in nearby Holyoke, but house their team offices at 100 Congress Street in Springfield.


title: Springfield, Massachusetts

text: Springfield has an official roller derby team: Pair O' Dice City Roller Derby. They are a non-profit organization which uses their roller derby games as fundraisers for groups such as Dakin Animal Shelter and the Shriners.


title: Springfield, Massachusetts

text: In 2010, Springfield was cited as the fourth "Greenest City" in the United States—the largest city cited in the Top 10. The recognition noted Springfield's numerous parks, the purity of its drinking water, its regional recycling center, and organizations like ReStore Home Improvement Center, which salvages building materials. Springfield features over of parkland distributed among 35 urban parks, including the grand, Forest Park. Well-known parks include the following, among others:


title: Springfield, Massachusetts

text: Springfield employs a strong-mayor form of city government. Springfield's mayor is Domenic J. Sarno, who has been serving since 2008.


title: Springfield, Massachusetts

text: The city's governmental bureaucracy consists of 33 departments, which administer a wide array of municipal services, e.g. police, fire, public works, parks, public health, housing, economic development, and the Springfield Public School System, New England's 2nd largest public school system.


title: Springfield, Massachusetts

text: Springfield's legislative body is its City Council, which features a mix of eight ward representatives—even though the city has more than twice that many neighborhoods, resulting in several incongruous "wards"—and five at-large city representatives, several of whom have served for well over a decade.


title: Springfield, Massachusetts

text: In 2003, the City of Springfield was on the brink of financial default, and thus taken over by a Commonwealth-appointed Finance Control Board until 2009. Disbanded in June of that year, the Control Board made great strides stabilizing Springfield's finances. While Springfield has achieved balanced budgets since 2009, the city has not enlarged its tax base, and thus many of its public works projects—which have been in the pipeline for years, some even decades—remain unfinished (e.g., repairs to Springfield's landmark Campanile). The construction of MGM Springfield, which opened in 2018, fueled a number of projects in the years leading up to and after its opening, with an estimated $3 billion of new development and infrastructure spending materializing.


title: Springfield, Massachusetts

text: Building off of the work of the Control Board, the city's finances have remained stable under Mayor Domenic J. Sarno's (2008–present) despite the Great Recession and several natural and man made disasters: June 1, 2011, tornado Springfield Tornado, Hurricane Irene, a freak October snow storm (which in some ways was more damaging than the tornado), and a large gas explosion in the downtown area in 2012. The city has recovered, however receiving a bond upgrade from Standard and Poor's Investment Services and the GFOA's Distinguished Budget Award for six consecutive years.


title: Springfield, Massachusetts

text: Like every other municipality in Massachusetts, Springfield has no judicial branch itself. Rather, it uses the Springfield-based state courts, which include Springfield district court and Hampden County Superior Court, both of which are based in Springfield. The Federal District Court also regularly hears cases in Springfield—now in an architecturally award-winning building on State Street, constructed in 2009.


title: Springfield, Massachusetts

text: Springfield became a city on May 25, 1852, by decree of the Massachusetts Legislature, after a decade-long internal dispute that resulted in the partition of Chicopee from Springfield, and thus the loss of two fifths of the city's population.


title: Springfield, Massachusetts

text: Springfield, like all municipalities in Massachusetts, is subject to limited home rule municipal power. The current city charter, in effect since 1959, uses a "strong mayor" government with most power concentrated in the mayor, as in Boston and elsewhere. The mayor representing the city's executive branch presents the budget, appoints commissioners and department heads, and in general runs the city. The mayor is former City Councilor Domenic Sarno, elected November 6, 2007, by a margin of 52.54% to 47.18% against incumbent Charles Ryan. He took office in January 2008. In November 2009 and 2011, Sarno won reelection, albeit—in the latter case—with just 22% of eligible Springfield voters voting.


title: Springfield, Massachusetts

text: The Springfield City Council, consisting of thirteen members, is the city's legislative branch. Elected every odd numbered year, eight of its members are elected to represent "wards", which are made of (sometimes incongruous) groupings of Springfield neighborhoods, e.g. Springfield's ethnic North End neighborhoods—Memorial Square and Brightwood—share a ward with Metro Center, Springfield's downtown. Five city council members are elected at-large. The City Council passes the city's budget, holds hearings, creates departments and commissions, and amends zoning laws.


title: Springfield, Massachusetts

text: The mayor's office and city council chambers are located in city hall—part of the Municipal Group in Metro Center, Springfield. The Finance Control Board met there as well.


title: Springfield, Massachusetts

text: Springfield is heavily Democratic. In the 2020 US Presidential Election Democrat Joe Biden received 72.4 percent of the vote to Republican Donald Trump's 25.7 percent.


title: Springfield, Massachusetts

text: In the past, efforts have been made to provide each of the city's eight wards a seat in the city council, instead of the current at-large format. There would still be some at-large seats under this format. The primary argument for this has been that City Councilors live in only four of the city's wards. An initiative to change the composition failed to pass the City Council twice. In 2007 Mayor Charles V. Ryan and City Councilor Jose Tosado proposed a home-rule amendment that would expand the council to thirteen members adding four seats to the existing nine member at large system, but allocated between eight ward and five at large seats. This home-rule petition was adopted by the City Council 8–1, and was later passed by the State Senate and House and signed by the Governor. On election day, November 6, 2007, city residents voted overwhelmingly in favor of changing the City Council and School Committee. The ballot initiative that established a new council with five at-large seats and eight ward seats passed 3–1. On November 3, 2009, Springfield held its first ward elections in 50 years.


title: Springfield, Massachusetts

text: The State Representatives are Orlando Ramos, Carlos González, Bud Williams, and Angelo Puppolo. The State Senator is Adam Gomez.


title: Springfield, Massachusetts

text: Richard Neal represents United States House of Representatives, Massachusetts District 1, and Elizabeth Warren and Edward Markey represent Springfield in the Federal Senate.


title: Springfield, Massachusetts

text: The Springfield Plan for citizenship education drew national attention in the 1940s for its efforts to combat racism.


title: Springfield, Massachusetts

text: Springfield has the second-largest school district in Massachusetts and in New England. It operates 38 elementary schools, six high schools, six middle schools (6–8), and seven specialized schools. The main high schools in the city include the High School of Commerce, Springfield Central High School, Roger L Putnam Vocational-Technical High School, and the Springfield High School of Science and Technology, better known as Sci-Tech. There are also two charter secondary schools in the City of Springfield: SABIS International, which ranks among the top 5% of high schools nationally in academic quality, and the Hampden Charter School of Science.


title: Springfield, Massachusetts

text: The city is required under a 1970s court order to balance schools racially, which had necessitated busing. However, since then, the city and the school's population has shifted and many of the neighborhoods are more integrated, calling into question the need for busing at all.


title: Springfield, Massachusetts

text: In June 2015, Massachusetts Governor Charlie Baker announced $3.2 million in grants to three underperforming middle schools in Springfield.


title: Springfield, Massachusetts

text: The Roman Catholic Diocese of Springfield operated five Catholic elementary schools in the city, all of which were consolidated into a single entity, St. Michael's Academy, in the autumn of 2009. The non-denominational Pioneer Valley Christian Academy is located in the suburban Sixteen Acres neighborhood, educating K–12. Non-sectarian elementary schools within the City of Springfield include the Pioneer Valley Montessori School in Springfield's Sixteen Acres neighborhood and Orchard Children's Corner in suburban Indian Orchard, a Pre-Kindergarten, among others.


title: Springfield, Massachusetts

text: The diocese runs Pope Francis Preparatory School (previously Cathedral High School), which is the largest Catholic high school in Western Massachusetts. Two nonsectarian private schools are also located in Springfield: Commonwealth Academy located on the former campus of the MacDuffie School (which moved to Granby, Massachusetts, in 2011 after 130 years in Springfield), and teaches grades four through twelve, soon to enroll students in grades K–12; and the Academy Hill School, which teaches kindergarten through grade eight.


title: Springfield, Massachusetts

text: Within of Springfield are many private prep schools, which can serve as day schools for Springfield students; they include the Williston Northampton School in Easthampton, Massachusetts; Wilbraham & Monson Academy in Wilbraham, Massachusetts; and Suffield Academy in Suffield, Connecticut.


title: Springfield, Massachusetts

text: The Knowledge Corridor boasts the second-largest concentration of higher learning institutions in the United States, with 32 universities and liberal arts colleges and over 160,000 university students in Greater Hartford–Springfield. This includes two college consortia, the Five Colleges, and the Cooperating Colleges of Greater Springfield, whose member institutions often collaborate in greater outreach programs. Within of Springfield's Metro Center, there are 18 universities and liberal arts colleges, which enroll approximately 100,000 students.


title: Springfield, Massachusetts

text: As of 2015, Springfield attracts over 20,000 university students per year. Its universities and colleges include Western New England University; Springfield College, famous as the birthplace of the sport of basketball (1891) and the nation's first physical education class, (1912); American International College, founded to educate America's immigrant population, is notable as the inventor of the Model Congress program. UMass Amherst relocated its urban design center graduate program to Court Square in Metro Center.


title: Springfield, Massachusetts

text: Several of Greater Springfield's institutions rank among the most prestigious in the world. For example, Amherst College, north of Springfield, and Smith College, north of Springfield, consistently rank among America's top 10 liberal arts colleges. Mount Holyoke College—the United States' first women's college—consistently ranks among America's Top 15 colleges, and it is located only north of Springfield. Hampshire College is located north of Springfield. The 30,000-student University of Massachusetts Amherst is located north of Springfield. Approximately west of Springfield, across the Memorial Bridge in Westfield, is Westfield State University, founded by noted education reformer Horace Mann. Westfield was the first university in America to admit students without regard to sex, race, or economic status.


title: Springfield, Massachusetts

text: Just outside Springfield's northern city limits is Elms College, a Catholic college. Likewise, just below Springfield's southern city limit in Longmeadow is Bay Path University; both schools were once all-women but are now co-ed.


title: Springfield, Massachusetts

text: In 1968, following the Pentagon's controversial closing of the Springfield Armory, Springfielders founded Springfield Technical Community College on behind the Springfield Armory National Park. Springfield Technical Community College is the only polytechnic community college in Massachusetts, and was founded to continue Springfield's tradition of technical innovation.


title: Springfield, Massachusetts

text: Holyoke Community College, north of Springfield, offers more traditional community college programming in Greater Springfield, as well as instruction in the culinary arts.


title: Springfield, Massachusetts

text: Springfield's largest local newspaper is "The Republican", which has also previously been known as the "Springfield Union-News & Sunday Republican", from when it merged with the "Springfield Union". Smaller papers such as "The Reminder" and the "Valley Advocate" also serve Greater Springfield.


title: Springfield, Massachusetts

text: Other newspapers serve specific communities of interest, such as "El Pueblo Latino" and "El Sol Latino", which serve the Hispanic community, as well as "Unity First" and the "AfAm Point of View", both of which serve the African-American community, and "The Rainbow Times", which serves Springfield's LGBT community.


title: Springfield, Massachusetts

text: Springfield has a long history of broadcast television, including two of the oldest UHF television stations on the air today.


title: Springfield, Massachusetts

text: Springfield proper is serviced exclusively by Comcast cable. Springfield was formerly wired with a "dual plant" cable system from 1980 until 2001, requiring an A/B switch for each home to watch programming and complicating VCR recordings.


title: Springfield, Massachusetts

text: Springfield was home to the first commercially licensed radio station in the United States, and the oldest radio station of any kind in New England: WBZ, which broadcast live from Springfield's luxurious Hotel Kimball at 140 Chestnut Street (now the Kimball Towers Condominiums) before moving to Boston in 1931.


title: Springfield, Massachusetts

text: Springfield is called the "Crossroads of New England" because it is the major shipping nexus from New York City, Boston, Montreal and the Great Lakes (via Albany, New York). Much of the cargo heading from one of these places to another crosses through the City of Springfield. As a geographical trade center, Springfield has more advantages than just being equidistant to these other large trade centers—it sits beside the Connecticut River, is located near some of the most fertile farmland in the Northeast, and is served by numerous rail lines and Interstate Highways, including I-90 (Mass Pike) and I-91, which connect New Haven, Hartford, Holyoke, Northampton, and Vermont to Springfield. One of the few spurs of I-91 in Massachusetts, I-291, runs through Springfield, and provides a secondary connection between I-90 and I-91.


title: Springfield, Massachusetts

text: Springfield Union Station, originally opened in 1926 and re-opened in 2017, is served by five Amtrak intercity routes: the "Vermonter", which runs from Washington, D.C. to St. Albans, Vermont; the "Lake Shore Limited", which runs from Chicago to Boston; the "Hartford Line", which runs from Springfield to New Haven; the "Valley Flyer", which runs from New Haven to Greenfield; and the "Northeast Regional", which runs from Springfield to D.C./Virginia. Amtrak relocated its operations into Union Station proper from their previous track side station building in June 2019. A high level train platform is under construction to improve the experience for rail passengers and is scheduled to be completed before the end of 2019.


title: Springfield, Massachusetts

text: CTRail's Hartford Line started operating from Union Station in June 2018, with Springfield as the northern terminus. Trains operate to New Haven, CT with multiple stops in Connecticut along the way. The line shares the same route and station listing as the Amtrak route of the same name, and the two are operated in conjunction as a commuter rail service for the region.


title: Springfield, Massachusetts

text: The New Haven–Springfield Line was upgraded in conjunction with the launch of the Hartford Line service. The project received funding from the Commonwealth of Massachusetts, the Federal Government, and the State of Connecticut. Amtrak trains on the route between New Haven and Springfield reach speeds of .


title: Springfield, Massachusetts

text: In 2011, Springfield Union Station started a $70 million renovation to become an intermodal transportation facility, allowing Peter Pan Bus, Greyhound Bus, and the PVTA to occupy a modern space next to the renovated Union Station. It was completed in June 2017.


title: Springfield, Massachusetts

text: There are no major freight yards in Springfield proper, but Connecticut Southern Railroad and CSX serve CSX's West Springfield yard across the Connecticut River.


title: Springfield, Massachusetts

text: The Pioneer Valley Transit Authority (PVTA) is based in Springfield and uses Union Station as its Springfield hub, next to the Gothic arch that denotes the entrance to downtown Springfield. The PVTA operates nineteen bus routes from Union Station. The PVTA's Springfield service area includes Springfield, Holyoke, Chicopee, Westfield, West Springfield, Ludlow, Agawam, and East Longmeadow. With transfers, it is possible to travel into PVTA's northern service area, which includes Northampton, Amherst, and Easthampton.


title: Springfield, Massachusetts

text: Intercity bus service is provided by Peter Pan Bus Lines and Greyhound Lines, both of which operate from Union Station. They provide service to destinations across the northeast United States.


title: Springfield, Massachusetts

text: Springfield is primarily served by Bradley International Airport, in Windsor Locks, Connecticut, south of downtown. It features over 100 daily departures to 30 destinations on nine airlines. It is also the primary airport for Hartford.


title: Springfield, Massachusetts

text: Taken in its entirety, Springfield has a moderate Walk Score of 59, however walkability varies between neighborhoods. Whereas the suburban neighborhood of Sixteen Acres is largely car-dependent with a score of 30, and Indian Orchard has a somewhat walkable rating of 54, the Metro Center area, with its grid central to stores, residences, and businesses, yields a Walk Score of 82.


title: Springfield, Massachusetts

text: The city's Connecticut River Walk Park offers a largely uninterrupted bike route from the North to South End, and is part of a broader plan by the Pioneer Valley Planning Commission to create a continuous route alongside the Connecticut River from Agawam to Holyoke, with construction and planning ongoing.


title: Springfield, Massachusetts

text: The city recently partnered with Bewegen (an e-bike share system provider) to install e-bike share stations in both Springfield and West Springfield.


title: Springfield, Massachusetts

text: The Springfield Water and Sewer Commission (created in its current form in 1996) owns several reservoirs and aqueducts, as well as hydropower and sewage treatment stations. The city purchased the Springfield Aqueduct Company in 1872.


title: Springfield, Massachusetts

text: Borden Brook Reservoir, located in the rural western Hampden County town of Blandford was completed in 1910. It feeds into the Cobble Mountain Reservoir (completed in 1931) located at the junction of the towns of Blandford, Granville and Russell. The Wild Cat Aqueduct carries water from the Cobble Mountain Reservoir to a hydroelectric generating station on the Granville–Russell border, at the Little River. Drinking water flows to the West Parish Water Filtration Plant in Westfield, and is then pumped to holding tanks at the top of Provin Mountain in Agawam.


title: Springfield, Massachusetts

text: The 1875 Ludlow Reservoir, also known as Springfield Reservoir, is maintained as an emergency water supply; it is located in Ludlow and fed via the Broad Brook Canal.


title: Springfield, Massachusetts

text: SWSC provides retail water in Springfield and Ludlow; wholesale water to Agawam, East Longmeadow, and Longmeadow; partial or peak service to Southwick, Westfield, and West Springfield; and emergency service to Chicopee and Wilbraham.


title: Springfield, Massachusetts

text: The Springfield Fire Department provides fire protection and emergency medical services to the city, and is one of the oldest fire departments in the United States.


title: Athens International Airport

text: Athens International Airport "Eleftherios Venizelos" (, "Diethnís Aeroliménas Athinón "Elefthérios Venizélos""), commonly initialised as AIA , is the largest international airport in Greece, serving the city of Athens and region of Attica. It began operation on 28 March 2001 (in time for the 2004 Summer Olympics) and is the main base of Aegean Airlines, as well as other smaller Greek airlines. It replaced the old Ellinikon International Airport. Athens International is currently a member of Group 1 of Airports Council International (over 25 million passengers) as of 2021, it is the 15th-busiest airport in Europe and the busiest and largest in the Balkans.


title: Athens International Airport

text: AIA is located between the towns of Markopoulo, Koropi, Spata and Loutsa, about to the east of central Athens ( by road, due to intervening hills). The airport is named after Elefthérios Venizélos, the prominent Cretan political figure and Prime Minister of Greece, who made a significant contribution to the development of Greek aviation and the Hellenic Air Force in the 1930s. As to-date, the airport is operated by AIA S.A. and ownership is divided between the Hellenic Republic (Greek State) and Private Sector in a 55%-45% stake following a PPP scheme for the airport company. Currently, private investors include the Copelouzos Group (5%) and PSP Investments of Canada (40%), following purchase of Hochtief's shares.