Brood parasitism is a form of parasitism

Parasitism is a type of non mutual relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host (e.g. Taenia solium). These are now called macroparasites (typically protozoa and helminths). The word parasite now also refers to microparasites, which are typically smaller, such as viruses and bacteria, and can be directly transmitted between hosts of the same species ^[1]. Examples of parasites include the plants mistletoe and cuscuta, and organisms such as leeches.

Unlike predators, parasites are generally much smaller than their host; both are special cases of consumer-resource interactions.^[2] Parasites show a high degree of specialization, and reproduce at a faster rate than their hosts. Classic examples of parasitism include interactions between vertebrate hosts and diverse animals such as tapeworms, flukes, the Plasmodium species, and fleas.

Parasitism is differentiated from the parasitoid relationship, though not sharply, by the fact that parasitoids generally kill or sterilise their hosts. Parasitoidism occurs in much the same variety of organisms that parasitism does.

The harm and benefit in parasitic interactions concern the biological fitness of the organisms involved. Parasites reduce host fitness in many ways, ranging from general or specialized pathology, such as parasitic castration and impairment of secondary sex characteristics, to the modification of host behaviour. Parasites increase their fitness by exploiting hosts for resources necessary for the parasite's survival, e.g. food, water, heat, habitat, and genetic dispersion.

Although the concept of parasitism applies unambiguously to many cases in nature, it is best considered part of a continuum of types of interactions between species, rather than an exclusive category. Particular interactions between species may satisfy some but not all parts of the definition. In many cases, it is difficult to demonstrate that the host is harmed. In others, there may be no apparent specialization on the part of the parasite, or the interaction between the organisms may be short-lived.

Etymology[link]

First used in English 1539, the word parasite comes from the Medieval French parasite, from the Latin parasitus, the latinisation of the Greek παράσιτος (parasitos), "one who eats at the table of another"^[3] and that from παρά (para), "beside, by"^[4] + σῖτος (sitos), "wheat".^[5] Coined in English in 1611, the word parasitism comes from the Greek παρά (para) + σιτισμός (sitismos) "feeding, fattening".^[6]

Types[link]

Parasites are classified based on their interactions with their hosts and on their life cycles.

Parasites that live on the surface of the host are called ectoparasites (e.g. some mites). Those that live inside the host are called endoparasites (including all parasitic worms). Endoparasites can exist in one of two forms: intercellular parasites (inhabiting spaces in the host’s body) or intracellular parasites (inhabiting cells in the host’s body). Intracellular parasites, such as protozoa, bacteria or viruses, tend to rely on a third organism, which is generally known as the carrier or vector.^{[citation needed]} The vector does the job of transmitting them to the host. An example of this interaction is the transmission of malaria, caused by a protozoan of the genus Plasmodium, to humans by the bite of an anopheline mosquito. Those parasites living in an intermediate position, being half-ectoparasites and half-endoparasites, are sometimes called mesoparasite.

An epiparasite is one that feeds on another parasite. This relationship is also sometimes referred to as hyperparasitism, exemplified by a protozoan (the hyperparasite) living in the digestive tract of a flea living on a dog.

Social parasites take advantage of interactions between members of social organisms such as ants or termites. In kleptoparasitism, parasites appropriate food gathered by the host. An example is the brood parasitism practiced by many species of cuckoo and cowbird, which do not build nests of their own but rather deposit their eggs in nests of other species and abandon them there. The host behaves as a "babysitter" as they raise the young as their own. If the host removes the cuckoo's eggs, some cuckoos will return and attack the nest to compel host birds to remain subject to this parasitism.^[7] The cowbird’s parasitism does not necessarily harm its host’s brood; however, the cuckoo may remove one or more host eggs to avoid detection, and furthermore the young cuckoo may heave the host’s eggs and nestlings out of the nest.

Parasitism can take the form of isolated cheating or exploitation among more generalized mutualistic interactions. For example, broad classes of plants and fungi exchange carbon and nutrients in common mutualistic mycorrhizal relationships; however, some plant species known as myco-heterotrophs "cheat" by taking carbon from a fungus rather than donating it.

Parasitoids are organisms whose larval development occurs inside or on the surface of another organism, resulting in the death of the host.^[8] This means that the interaction between the parasitoid and the host is fundamentally different from that of a true parasite and shares some of the characteristics of predation.

An adelpho-parasite is a parasite in which the host species is closely related to the parasite, often being a member of the same family or genus. An example of this is the citrus blackfly parasitoid, Encarsia perplexa, unmated females of which may lay haploid eggs in the fully developed larvae of their own species. These result in the production of male offspring.^[9] The marine worm Bonellia viridis has a similar reproductive strategy, although the larvae are planktonic.^[10]

Host defenses[link]

This section requires expansion.

Hosts respond to parasitisms in many ways ranging from the morphological to the behavioural. In some cases, plants produce toxins to deter parasitic fungi and bacteria.^{[citation needed]} Vertebrate animals have developed complex immune systems, which can target parasites through contact with bodily fluids.^{[citation needed]} Animals are also known to resort to behavioral defenses, examples of which are the avoidance by sheep of open pastures during spring, when roundworm eggs accumulated over the previous year hatch en masse;^{[citation needed]} and the ingestion of alcohol by infected fruit flies as self medication against blood-borne parasites.^[11] In humans, parasite immunity is developed prominently by Immunoglobulin E antibodies.

Evolutionary aspects[link]

Biotrophic parasitism is a common mode of life that has arisen independently many times in the course of evolution. Depending on the definition used, as many as half of all animals have at least one parasitic phase in their life cycles,^[12] and it is also frequent in plants and fungi. Moreover, almost all free-living animals are host to one or more parasites taxa.^[12]

Restoration of a Tyrannosaurus with parasite infections. A 2009 study showed that holes in the skulls of several specimens might have been caused by Trichomonas-like parasites^[13]

Parasites evolve in response to the defense mechanisms of their hosts. As a result of host defenses, some parasites evolve adaptations that are specific to a particular host taxon, specializing to the point where they infect only a single species. Such narrow host specificity can be costly over evolutionary time, however, if the host species becomes extinct. Therefore many parasites can infect a variety of more or less closely related host species, with different success rates.

Host defenses also evolve in response to attacks by parasites. Theoretically, parasites may have an advantage in this evolutionary arms race because of their more rapid generation time. Hosts reproduce less quickly than parasites, and therefore have fewer chances to adapt than their parasites do over a given span of time.

In some cases, a parasite species may coevolve with its host taxa. Long-term coevolution sometimes leads to a relatively stable relationship tending to commensalism or mutualism, as, all else being equal, it is in the evolutionary interest of the parasite that its host thrives. A parasite may evolve to become less harmful for its host or a host may evolve to cope with the unavoidable presence of a parasite-- to the point that the parasite's absence causes the host harm. For example, although animals infected with parasitic worms are often clearly harmed, and therefore parasitized, such infections may also reduce the prevalence and effects of autoimmune disorders in animal hosts, including humans.^[14]

Competition between parasites tends to favor faster reproducing and therefor more virulent parasites. Parasites whose life cycle involves the death of the host, to exit the present host and sometimes to enter the next, evolve to be more virulent or even alter the behavior or other properties of the host to make it more vulnerable to predators. Parasites that reproduce largely to the offspring of the previous host tend to become less virulent or mutualist, so that its hosts reproduce more effectively.^[1]

The presumption of a shared evolutionary history between parasites and hosts can sometimes elucidate how host taxa are related. For instance, there has been dispute about whether flamingos are more closely related to the storks and their allies, or to ducks, geese and their relatives. The fact that flamingos share parasites with ducks and geese is evidence these groups may be more closely related to each other than either is to storks.

Parasitism is part of one explanation for the evolution of secondary sex characteristics seen in breeding males throughout the animal world, such as the plumage of male peacocks and manes of male lions. According to this theory, female hosts select males for breeding based on such characteristics because they indicate resistance to parasites and other disease.

Co-speciation[link]

In rare cases, a parasite may even undergo co-speciation with its host. One particularly remarkable example of co-speciation exists between the simian foamy virus (SFV) and its primate hosts. In one study, the phylogenies of SFV polymerase and the mitochondrial cytochrome oxidase subunit II from African and Asian primates were compared.^[15] Surprisingly, the phylogenetic trees were very congruent in branching order and divergence times. Thus, the simian foamy viruses may have co-speciated with Old World primates for at least 30 million years.

Ecology[link]

Quantitative ecology[link]

A single parasite species usually has an aggregated distribution across host individuals, which means that most hosts harbor few parasites, while a few hosts carry the vast majority of parasite individuals. This poses considerable problems for students of parasite ecology: the use of parametric statistics should be avoided.^{[citation needed]} Log-transformation of data before the application of parametric test, or the use of non-parametric statistics is recommended by several authors. However, this can give rise to further problems.^[16] Therefore, modern day quantitative parasitology is based on more advanced biostatistical methods.

Diversity ecology[link]

Hosts represent discrete habitat patches that can be occupied by parasites. A hierarchical set of terminology has come into use to describe parasite assemblages at different host scales.

Infrapopulation

All the parasites of one species in a single individual host.

Metapopulation

All the parasites of one species in a host population.

Infracommunity

All the parasites of all species in a single individual host.

Component community

All the parasites of all species in a host population.

Compound community

All the parasites of all species in all host species in an ecosystem.

The diversity ecology of parasites differs markedly from that of free-living organisms. For free-living organisms, diversity ecology features many strong conceptual frameworks including Robert MacArthur and E. O. Wilson's theory of island biogeography, Jared Diamond's assembly rules and, more recently, null models such as Stephen Hubbell's unified neutral theory of biodiversity and biogeography. Frameworks are not so well-developed for parasites and in many ways they do not fit the free-living models. For example, island biogeography is predicated on fixed spatial relationships between habitat patches ("sinks"), usually with reference to a mainland ("source"). Parasites inhabit hosts, which represent mobile habitat patches with dynamic spatial relationships. There is no true "mainland" other than the sum of hosts (host population), so parasite component communities in host populations are metacommunities.

Nonetheless, different types of parasite assemblages have been recognized in host individuals and populations, and many of the patterns observed for free-living organisms are also pervasive among parasite assemblages. The most prominent of these is the interactive-isolationist continuum. This proposes that parasite assemblages occur along a cline from interactive communities, where niches are saturated and interspecific competition is high, to isolationist communities, where there are many vacant niches and interspecific interaction is not as important as stochastic factors in providing structure to the community. Whether this is so, or whether community patterns simply reflect the sum of underlying species distributions (no real "structure" to the community), has not yet been established.

Adaptation[link]

Parasites infect hosts that exist within their same geographical area (sympatric) more effectively. This phenomenon supports the "Red Queen hypothesis—which states that interactions between species (such as host and parasites) lead to constant natural selection for adaptation and counter adaptation."^[17] The parasites track the locally common host phenotypes, therefore the parasites are less infective to allopatric (from different geographical region) hosts.^{[citation needed]}

Experiments published in 2000 discuss the analysis of two different snail populations from two different sources- Lake Ianthe and Lake Poerua in New Zealand. The populations were exposed to two pure parasites (digenetic trematode) taken from the same lakes. In the experiment, the snails were infected by their sympatric parasites, allopatric parasites and mixed sources of parasites. The results suggest that the parasites were more highly effective in infecting their sympatric snails than their allopatric snails. Though the allopatric snails were still infected by the parasites, the infectivity was much less when compared to the sympatric snails. Hence, the parasites were found to have adapted to infecting local populations of snails.^[17]

Transmission[link]

Life cycle of Entamoeba histolytica, an anaerobic parasitic protozoan.

Parasites inhabit living organisms and therefore face problems that free-living organisms do not. Hosts, the only habitats in which parasites can survive, actively try to avoid, repel, and destroy parasites. Parasites employ numerous strategies for getting from one host to another, a process sometimes referred to as parasite transmission or colonization.

Some endoparasites infect their host by penetrating its external surface, while others must be ingested. Once inside the host, adult endoparasites need to shed offspring into the external environment to infect other hosts. Many adult endoparasites reside in the host’s gastrointestinal tract, where offspring can be shed along with host excreta. Adult stages of tapeworms, thorny-headed worms and most flukes use this method.

Among protozoan endoparasites, such as the malarial parasites and trypanosomes, infective stages in the host’s blood are transported to new hosts by biting-insects, or vectors.

Larval stages of endoparasites often infect sites in the host other than the blood or gastrointestinal tract. In many such cases, larval endoparasites require their host to be consumed by the next host in the parasite’s life cycle in order to survive and reproduce. Alternatively, larval endoparasites may shed free-living transmission stages that migrate through the host’s tissue into the external environment, where they actively search for or await ingestion by other hosts. The foregoing strategies are used, variously, by larval stages of tapeworms, thorny-headed worms, flukes and parasitic roundworms.

Some ectoparasites, such as monogenean worms, rely on direct contact between hosts. Ectoparasitic arthropods may rely on host-host contact (e.g. many lice), shed eggs that survive off the host (e.g. fleas), or wait in the external environment for an encounter with a host (e.g. ticks). Some aquatic leeches locate hosts by sensing movement and only attach when certain temperature and chemical cues are present.

Some parasites modify host behavior to make transmission to other hosts more likely. For example, in California salt marshes, the fluke Euhaplorchis californiensis reduces the ability of its killifish host to avoid predators.^[18] This parasite matures in egrets, which are more likely to feed on infected killifish than on uninfected fish. Another example is the protozoan Toxoplasma gondii, a parasite that matures in cats but can be carried by many other mammals. Uninfected rats avoid cat odors, but rats infected with T. gondii are drawn to this scent, which may increase transmission to feline hosts.^[19]

Roles in ecosystems[link]

Modifying the behavior of infected hosts, to make transmission to other hosts more likely to occur, is one way parasites can affect the structure of ecosystems. For example, in the case of Euhaplorchis californiensis (discussed above) it is plausible that the local predator and prey species might be different if this parasite were absent from the system.

Although parasites are often omitted in depictions of food webs, they usually occupy the top position. Parasites can function like keystone species, reducing the dominance of superior competitors and allowing competing species to co-exist.

Many parasites require multiple hosts of different species to complete their life cycles and rely on predator-prey or other stable ecological interactions to get from one host to another. In this sense, the parasites in an ecosystem reflect the "health" of that system.

Effects on Host Behavior[link]

Animal behavior is typically motivated by proximate level mechanisms that promote particular actions. In most cases, proximate level mechanisms are a result of the individual’s interaction with the environment but some parasites have been shown to interact and manipulate these mechanisms to produce behaviors beneficial to the parasite alone. Parasites are known to drastically affect how animals behave. ^[20] Parasites most commonly target the central nervous system (CNS) in order to alter animal behavior. By affecting hormone secretions or by physical restructuring, parasites successfully change how an animal’s body functions and delivers, interprets and and reacts to messages. Some parasites, like toxoplasma, form vacuoles that travel through the nervous system interrupting key functions in intraneural communications. The emerald jewel wasp alters behavior through the injection of venom directly into the host’s brain, causing hypokinesia. ^[21] Parasitic life cycles can have the capacity to infect a singular host or a series of hosts. Direct life cycles are related to a single host while indirect life cycles, or complex life cycles, rely on a series of intermediate hosts in order to complete the life cycle. Indirect life cycles likely arose in order to increase the efficiency of spreading to new hosts. By using intermediary’s the parasite DNA can spread to several hosts instead of remain concentrated within a single host, therefore promising a higher likelihood of DNA continuation. In some cases, intermediary hosts are accidental. ^[22] Mud-snails (corophium volutator) from Atlantic mud flats have been observed to carry cyst like infections, caused by trematoad, specifically gynaecotyla adunca, infection. The trematoads enter the host body and create cyst formations over the snail’s body. As the infection progresses, the mud-snails are more apt to crawling behavior during daylight hours. Crawling, instead of burrowing, increases the snails risk to shorebird predation, which is the trematoads primary host. A side-effect of the increased crawling behavior is an increased likelihood of reproduction. Crawling is the mud-snails primary means to interaction with mates. Further, infected mud-snails appear to have a higher reproductive rate than uninfected, suggesting a host adaptation to the parasitic infection. ^[23] Toxoplasma gondii is a noted case of unintended intermediary hosts. Typically, toxoplasma resides in animals from the felidae family, exiting through excretion in order to infect rodents. After infecting the rodents, who consume the felidae fecal matter simultaneously consuming the toxoplasma, they begin to alter the rodent’s behavior. Rodents become more extroverted and less fearful of felines. Toxoplasma, however, has recently made a host of humans. In doing so, human behavior has been altered in similar ways to rodents. Further, toxoplasma has been linked to cases of schizophrenia. ^[24]

The emerald jewel wasp (Ampulex compressa) takes advantage of its host, the American cockroach (Periplaneta americana) specifically as a food source and home for its growing larvae. The wasp begins by injecting venom into the brain, specifically the central nervous system system of the cockroach paralyzing it by putting it into a state of “hypokinesia”. “Hypokinesia...is a reversible long-term lethargy characterized by lack of spontaneous movement or response to external stimuli,” (Banks and Adams). After dragging the cockroach to a burrow, the wasp deposits an egg into its carcass, burying it for the growing larvae to feed off of until it emerges in 6 weeks, leaving nothing but a hard outer cockroach shell. ^[25] Although the circuitry in control of movement is functional, the nervous system acts from a depressed state. They are not killed by this hypokinesia, but would recover if not for the larvae eating them from the inside out. The movement is controlled by dopamine and octopamine which affect transmission of interneurons involved in the natural response to escape. Reduced motor activity results from a reduction of these amines. ^[26]

Value[link]

Although parasites are generally considered to be harmful, the eradication of all parasites would not necessarily be a noble aim. Among other things, parasites may account for as much as or more than half of life's diversity; they perform an important ecological role (by weakening prey) that ecosystems would take some time to adapt to; and without parasites organisms may eventually tend to asexual reproduction, dinimishing the diversity of sexually dimorphic traits.^[27] Parasites provide an opportunity for the transfer of genetic material between species. On rare but significant occasions this may provide a path for evolutionary changes that would not otherwise occur, or that would otherwise take even longer.^[1]

See also[link]

References[link]

Further reading[link]

• Zimmer, Carl (2001). Parasite Rex. Free Press. pp. 320. ISBN 0-7432-0011-X. 
• Combes, Claude (2005). The Art of Being a Parasite. The University of Chicago Press. pp. 280. ISBN 0-226-11438-4. 
• Desowitz, Robert (1998). Who Gave Pinta to the Santa Maria?. Harvest Books. pp. 264. ISBN 0-15-600585-9. 

External links[link]

Wikimedia Commons has media related to: Parasitism

• Parasitism—Knol
• Toxoplasmosis
• Parasitology Parasites Zoonoses—(Polish/English) over 50 movies (Filmoteka) and over 250 photos (Fotogaleria/Photogallery) with human and animal parasites.
• Aberystwyth University: Parasitology—class outline with links to full text articles on parasitism and parasitology.
• KSU: Parasitology Research—parasitology articles and links.
• Medical Parasitology—online textbook.
• Division of Parasitic Diseases, Centers for Disease Control and Prevention
• VCU Virtual Parasite Project—Virtual Parasite Project at Virginia Commonwealth University's Center for the Study of Biological Complexity
• Parasites World—Parasites articles and links.
• Parasitic and Parasitoid Alien Species in Science Fiction Movies

Nicholas Rodney "Nick" Drake (19 June 1948 – 25 November 1974) was an English singer-songwriter and musician who was known for his sombre guitar-based songs. He failed to find a wide audience during his lifetime but his work has gradually achieved wider notice and recognition.^[2][3][4] Drake signed to Island Records when he was 20 years old and released his debut album, Five Leaves Left, in 1969. By 1972, he had recorded two more albums—Bryter Layter and Pink Moon. Neither sold more than 5,000 copies on initial release.^[5] Drake's reluctance to perform live, or be interviewed, contributed to his lack of commercial success.

Drake suffered from depression and insomnia throughout his life and these topics were often reflected in his lyrics. On completion of his third album, 1972's Pink Moon, he withdrew from both live performance and recording, retreating to his parents' home in rural Warwickshire. There is no known footage of the adult Drake; he was only ever captured in still photographs and in home footage from his childhood.^[6] On 25 November 1974, Drake died from an overdose of amitriptyline, a prescribed antidepressant; he was 26 years old.

Drake's music remained available through the mid-1970s, but the 1979 release of the retrospective album Fruit Tree caused his back catalogue to be reassessed. By the mid-1980s Drake was being credited as an influence by such artists as Robert Smith, David Sylvian and Peter Buck. In 1985, The Dream Academy reached the UK and US charts with "Life in a Northern Town", a song written for and dedicated to Drake.^[7] By the early 1990s, he had come to represent a certain type of "doomed romantic" musician in the UK music press and was frequently cited as an influence by artists including Kate Bush, Paul Weller and The Black Crowes.^[8] His first biography appeared in 1997, and was followed in 1998 by the documentary film A Stranger Among Us. In 2000, Volkswagen featured the title track from Pink Moon in a television advertisement and, within a month, more Drake records were sold than in the previous 30 years.

Early life[link]

His father, Rodney Drake (1908–1988), had moved to Rangoon, Burma, in the early 1930s to work as an engineer with the Bombay Burmah Trading Corporation.^[9] There, in 1934, his father met the daughter of a senior member of the Indian Civil Service, Mary Lloyd (1916–1993), known to her family as "Molly". Rodney Drake proposed to Molly in 1936, though the couple had to wait a year until she turned 21 before her family allowed them to marry.^[10] In 1950, they returned to Warwickshire^[11] to live in the country, at a house named Far Leys, in the village of Tanworth-in-Arden in west Warwickshire, just south of Solihull. Drake had one older sister, Gabrielle, later a successful film and television actress. Both parents were musically inclined and they each wrote pieces of music. Recordings of Molly Drake's songs, which have come to light following her, death are remarkably similar in tone and outlook to the later work of her son.^[12] Mother and son shared a similar fragile vocal delivery and both Gabrielle and biographer Trevor Dann have noted a parallel sense of foreboding and fatalism in their music.^[12][13] Encouraged by his mother, Drake learned to play piano at an early age and began to compose his own songs which he recorded on a reel-to-reel tape recorder she kept in the family drawing room.^[4]

In 1957, Drake enrolled at Eagle House School, an English preparatory boarding school in Berkshire. Five years later, he went on to public school at Marlborough College in Wiltshire, where his father, grandfather, and great-grandfather had all attended. He developed an interest in sport, becoming an accomplished sprinter (his record for the 100-yard dash still stands)^[14] and captain of the school's rugby team for a time. He was also Head of House in C1, the College's largest house. School friends recall Drake at this time as having been confident and "quietly authoritative", while often aloof in his manner.^[15] His father Rodney remembered, "In one of his reports [the headmaster] said that none of us seemed to know him very well. All the way through with Nick. People didn't know him very much."^[16]

Drake played piano in the school orchestra, and learned clarinet and saxophone. He formed a band, The Perfumed Gardeners, with four schoolmates in 1964 or 1965. With Drake on piano and occasional alto sax and vocals, the group performed Pye covers and jazz standards, as well as Yardbirds and Manfred Mann numbers. The line-up briefly included Chris de Burgh, but he was soon ejected as his taste was seen as "too poppy" by the other members.^[17] Drake's academic performance began to deteriorate, and while he had accelerated a year in Eagle House, at Marlborough he began to neglect his studies in favour of music. He attained seven GCE O-Levels in 1963, but this was fewer than his teachers had been expecting, and he failed "Physics with Chemistry".^[18] In 1965, Drake paid £13 for his first acoustic guitar, and was soon experimenting with open tuning and finger-picking techniques.^[14]

In 1966, Drake won a scholarship to study English literature at Fitzwilliam College, University of Cambridge. He delayed attendance to spend six months at the University of Aix-Marseille, France, beginning in February 1967. While in Aix, he began to practise guitar in earnest, and to earn money would often busk with friends in the town centre. Drake began to smoke cannabis, and that spring he travelled with friends to Morocco, because, according to travelling companion Richard Charkin, "that was where you got the best pot".^[19] Drake most likely began using LSD while in Aix,^[20] and lyrics written during this period—in particular for the song "Clothes of Sand"—are suggestive of an interest in hallucinogens.^[21]

Cambridge[link]

Upon returning to England, he moved into his sister's flat in Hampstead, before enrolling at Cambridge that October. His tutors found him to be a bright student, but unenthusiastic and unwilling to apply himself to study.^[22] Dann notes that he had difficulty connecting with staff and fellow students alike, and points out that official matriculation photographs from this time reveal a sullen and unimpressed young man.^[23] Cambridge placed much emphasis on its rugby and cricket teams, yet by this time Drake had lost interest in playing sport, preferring to stay in his college room smoking marijuana, and listening to and playing music. According to fellow student (now psychiatrist) Brian Wells: "they were the rugger buggers and we were the cool people smoking dope."^[23] In September 1967, he met Robert Kirby, a music student who went on to orchestrate many of the string and woodwind arrangements for Drake's first two albums.^[24] By this time, Drake had discovered the British and American folk music scenes, and was influenced by performers such as Bob Dylan, Josh White and Phil Ochs. He began performing in local clubs and coffee houses around London, and in February 1968, while playing support to Country Joe and the Fish at the Roundhouse in Camden Town, made an impression on Ashley Hutchings, bass player with Fairport Convention.^[25] Hutchings recalls being impressed by Drake's skill as a guitarist, but even more so by "the image. He looked like a star. He looked wonderful, he seemed to be 7 ft."^[16]

Hutchings introduced Drake to the 25-year-old American producer Joe Boyd, owner of the production and management company Witchseason Productions. The company was, at the time, licensed to Island Records,^[26] and Boyd, as the man who had discovered Fairport Convention and been responsible for introducing John Martyn and The Incredible String Band to a mainstream audience, was a significant and respected figure on the UK folk scene.^[16] He and Drake formed an immediate bond, and the producer acted as a mentor figure to Drake throughout his career. A four-track demo, recorded in Drake's college room in the spring of 1968, led Boyd to offer a management, publishing, and production contract to the 20-year-old, and to initiate work on a debut album. According to Boyd:

In those days you didn't have cassettes—he brought a reel-to-reel tape [to me] that he'd done at home. Half way through the first song, I felt this was pretty special. And I called him up, and he came back in, and we talked, and I just said, "I'd like to make a record." He stammered, "Oh, well, yeah. Okay." Nick was a man of few words.^[16]

In a 2004 interview, Drake's friend Paul Wheeler remembered the excitement caused by his seeming big break, and recalled that the singer had already decided not to complete his third year at Cambridge.^[16]

Career[link]

[edit] Five Leaves Left

Drake began recording his debut album Five Leaves Left later in 1968, with Boyd assuming the role of producer. The sessions took place in Sound Techniques studio, London, with Drake skipping lectures to travel by train to the capital. Inspired by John Simon's production of Leonard Cohen's first album, Boyd was keen that Drake's voice would be recorded in a similar close and intimate style, "with no shiny pop reverb".^[27] He also sought to include a string arrangement similar to Simon's, "without overwhelming... or sounding cheesy".^[27] To provide backing, Boyd enlisted various contacts from the London folk rock scene, including Fairport Convention guitarist Richard Thompson and Pentangle bassist Danny Thompson (no relation).^[28] He recruited John Wood as engineer, and drafted Richard Hewson in to provide the string arrangements.

Initial recordings did not go well: the sessions were irregular and rushed, taking place during studio downtime borrowed from Fairport Convention's production of their Unhalfbricking album. Tension arose between artist and producer as to the direction the album should take: Boyd was an advocate of George Martin's "using the studio as an instrument" approach, while Drake preferred a more organic sound. Dann has observed that Drake appears "tight and anxious" on bootleg recordings taken from the sessions, and notes a number of Boyd's unsuccessful attempts at instrumentation.^[29] Both were unhappy with Hewson's contribution, which they felt was too mainstream in sound for Drake's songs.^[30] Drake suggested using his college friend Robert Kirby as a replacement, although Boyd was sceptical at taking on an amateur music student lacking prior recording experience. However, he was impressed by Drake's uncharacteristic assertiveness, and agreed to a trial.^[31] Kirby had previously presented Drake with some arrangements for his songs,^[26] and went on to provide a spare chamber music quartet score associated with the sound of the final album.^[32] However, Kirby did not feel confident enough to score the album's centerpiece "River Man", and Boyd was forced to stretch the Witchseason budget to hire the veteran composer Harry Robinson, with the instruction that he echo the tone of Delius and Ravel.

	"River Man" noicon "River Man" is noted for its 5/4 time, harmonic changes and use of prosody. An early solo acoustic version recorded in Drake's Cambridge college bedroom appears on the 2004 compilation Made To Love Magic.[33]
Problems listening to this file? See media help.

Post-production difficulties led to the release being delayed by several months. It has been alleged that the album was poorly marketed and supported,^[34] though the inclusion of the opening track "Time Has Told Me" on the Island Records sampler Nice Enough to Eat brought him a very wide audience (a track from his second album was likewise included on the subsequent sampler Bumpers). Drake was featured in full-page interviews in the pop press. In July, Melody Maker referred to the album as "poetic" and "interesting", though NME wrote in October that there was "not nearly enough variety to make it entertaining".^[35] It received radio plays from the BBC's more progressive disc-jockeys such as John Peel^[36] and Bob Harris. Drake was unhappy with the inlay sleeve, which printed songs in the wrong running order and reproduced verses omitted from the recorded versions.^[37] In an interview his sister Gabrielle said: "He was very secretive. I knew he was making an album but I didn't know what stage of completion it was at until he walked into my room and said, 'There you are.' He threw it onto the bed and walked out!"^[26]

[edit] Bryter Layter

Drake ended his studies at Cambridge nine months before graduation, and in autumn 1969 moved to London to concentrate on a career in music.^[38] His father remembered "writing him long letters, pointing out the disadvantages of going away from Cambridge ... a degree was a safety net, if you manage to get a degree, at least you have something to fall back on; his reply to that was that a safety net was the one thing he did not want."^[12] Drake spent his first few months in the capital drifting from place to place, occasionally staying at his sister's Kensington flat, but usually sleeping on friends’ sofas and floors.^[39] Eventually, in an attempt to bring some stability and a telephone into Drake's life, Boyd organised and paid for a ground floor bedsit in Belsize Park, Camden.^[40]

In August, Drake recorded three songs for the BBC's John Peel show. Two months later, he opened for Fairport Convention at the Royal Festival Hall in London, followed by appearances at folk clubs in Birmingham and Hull. Remembering the performance in Hull, folk singer Michael Chapman commented:

The folkies did not take to him; [they] wanted songs with choruses. They completely missed the point. He didn't say a word the entire evening. It was actually quite painful to watch. I don't know what the audience expected, I mean, they must have known they weren't going to get sea–shanties and sing-alongs at a Nick Drake gig!^[25]

The experience reinforced Drake's decision to retreat from live appearances; the few concerts he did play around this time were usually brief, awkward, and poorly attended. Drake seemed unwilling to perform and rarely addressed his audience. As many of his songs were played in different tunings, he frequently paused to retune between numbers.^[41]

	"Northern Sky" noicon "Northern Sky" features piano, organ and celesta performed by John Cale. Drake was reportedly in awe of Cale's musical ability.
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Although the publicity generated by Five Leaves Left was minor, Boyd was keen to build on what momentum there was. 1970's Bryter Layter, again produced by Boyd and engineered by Wood, introduced a more upbeat,^[42] jazzier^[43] sound. Disappointed by his debut's poor commercial performance, Drake sought to move away from his pastoral sound, and agreed to his producer's suggestions to include bass and drum tracks on the recordings. "It was more of a pop sound, I suppose", Boyd later said, "I imagined it as more commercial."^[44] Like its predecessor, the album featured musicians from Fairport Convention, as well as contributions from John Cale on two songs: "Northern Sky" and "Fly". Trevor Dann has noted that while sections of "Northern Sky" sound more characteristic of Cale, the song was the closest Drake came to a release with chart potential.^[45] In his 1999 autobiography, Cale admits to using heroin during this period,^[46] and his older friend Brian Wells began to suspect that Drake was also using.^[47] Both Boyd and Wood were confident that the album would be a commercial success,^[48] but it went on to sell fewer than 3,000 copies. Reviews were again mixed: while Record Mirror praised Drake as a "beautiful guitarist—clean and with perfect timing, [and] accompanied by soft, beautiful arrangements", Melody Maker described the album as "an awkward mix of folk and cocktail jazz".^[41]

Soon after the release, Boyd sold Witchseason to Island Records, and moved to Los Angeles to work with Warner Brothers in the development of soundtracks for film. The loss of this key mentor figure, coupled with the album's poor sales, led Drake to further retreat into depression. His attitude to London had changed: he was unhappy living alone, and visibly nervous and uncomfortable performing at a series of concerts in early 1970. In June, Drake gave one of his final live appearances at Ewell Technical College, London. Ralph McTell, who also performed that night, remembered that "Nick was monosyllabic. At that particular gig he was very shy. He did the first set and something awful must have happened. He was doing his song 'Fruit Tree' and walked off halfway through it. Just left the stage."^[49] His frustration turned to depression,^[50] and in 1971 Drake was persuaded by his family to visit a psychiatrist at St Thomas's Hospital, London. He was prescribed a course of antidepressants, but he felt uncomfortable and embarrassed about taking them, and tried to hide the fact from his friends.^[51] He knew enough about drugs to worry about their side effects, and was concerned about how they would react with his regular marijuana use.^[52]

[edit] Pink Moon

Island Records was keen that Drake promote Bryter Layter through press interviews, radio sessions and live appearances. Drake, who was by this time smoking what Kirby has described as "unbelievable amounts" of marijuana^[53] and exhibiting "the first signs of psychosis", refused. By the winter of 1970, he had isolated himself in London.^[38] Disappointed by the reaction to Bryter Layter, he turned his thoughts inwards, and withdrew from family and friends. He rarely left his flat, and then only to play an occasional concert or to buy drugs. "This was a very bad time", his sister Gabrielle Drake recalled, "He once said to me that everything started to go wrong from [this] time on, and I think that was when things started to go wrong."^[54]

Although Island neither expected nor wanted a third album,^[55] Drake approached Wood in October 1971 to begin work on what would be his final release. The sessions took place over two nights, with only Drake and Wood present in the studio.^[4] The bleak songs of Pink Moon are short, and the eleven-track album lasts only 28 minutes, a length described by Wood as "just about right. You really wouldn't want it to be any longer."^[16] Drake had expressed dissatisfaction with the sound of Bryter Layter, and believed that the string, brass and saxophone arrangements had resulted in a sound that was "too full, too elaborate".^[56] Drake appears on Pink Moon accompanied only by his own carefully recorded guitar save for a single piano overdub on the title track. "He was very determined to make this very stark, bare record," Wood later recalled. "He definitely wanted it to be him more than anything. And I think, in some ways, Pink Moon is probably more like Nick is than the other two records."^[57]

	"Pink Moon" noicon The piano overdub on "Pink Moon" displays a musicality absent from some of the barer tracks on Drake's final album.[58]
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Drake delivered the tapes of Pink Moon to Chris Blackwell at Island Records, contrary to a popular legend which claims he dropped them off at the receptionist's desk without saying a word.^[59] An advertisement for the album placed in Melody Maker in February opened with "Pink Moon—Nick Drake's latest album: the first we heard of it was when it was finished."^[60] Pink Moon went on to sell fewer copies than either of its predecessors, although it did receive some favourable reviews. In Zigzag magazine, Connor McKnight wrote, "Nick Drake is an artist who never fakes. The album makes no concession to the theory that music should be escapist. It's simply one musician's view of life at the time, and you can't ask for more than that."^[61]

Blackwell felt Pink Moon had the potential to bring Drake to a mainstream audience; however, his staff were disappointed by the artist's unwillingness to undertake any promotional activity. A&R manager Muff Winwood recalls "tearing his hair out" in frustration, and admits that without Blackwell's enthusiastic support, "the rest of us would have given him the boot."^[62] However, following persistent nagging from Boyd, Drake agreed to an interview with Jerry Gilbert of Sounds Magazine.^[63] The "shy and introverted folk singer" spoke of his dislike of live appearances and very little else.^[64] "There wasn't any connection whatsoever", Gilbert has said. "I don't think he made eye contact with me once. If you wanted to be uncharitable, you could say he was just a spoiled boy with a silver spoon and went around feeling sorry for himself."^[64] Disheartened and convinced he would be unable to write again, Drake decided to retire from music. He toyed with the idea of a different career, even considering the Army.^[65]

Final years[link]

In the months following Pink Moon's release, Drake became increasingly antisocial and distant from those close to him.^[66] He returned to live at his parents' home in Tanworth-in-Arden, and while he resented the regression, he accepted that his illness made it necessary. "I don't like it at home", he told his mother, "but I can't bear it anywhere else."^[12] His return was often difficult for his family; as his sister Gabrielle explained, "good days in my parents' home were good days for Nick, and bad days were bad days for Nick. And that was what their life revolved around, really."^[16]

	"Black Eyed Dog" noicon "Black Eyed Dog" from one of Drake's final recording sessions in February, 1974. The title was inspired by Winston Churchill's description, taken from Samuel Johnson, of depression as a black dog.[67]
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He lived a frugal existence, his only source of income being a £20-a-week retainer he received from Island Records. At one point he could not afford a new pair of shoes.^[68] He would often disappear for days, sometimes turning up unannounced at friends' houses, uncommunicative and withdrawn. Robert Kirby described a typical visit: "He would arrive and not talk, sit down, listen to music, have a smoke, have a drink, sleep there the night, and two or three days later he wasn't there, he'd be gone. And three months later he'd be back."^[69]

Referring to this period, John Martyn (who in 1973 wrote the title song of his album Solid Air for and about Drake) described him as the most withdrawn person he'd ever met.^[70] He would borrow his mother's car and drive for hours without purpose on occasion, until he ran out of petrol and had to ring his parents to ask to be collected. Friends have recalled the extent to which his appearance had changed.^[71] During particularly bleak periods of his illness, he refused to wash his hair or cut his nails.^[65] Early in 1972, Drake suffered a nervous breakdown, and was hospitalized for five weeks.^[47]

In February 1974, Drake again contacted John Wood, stating he was ready to begin work on a fourth album.^[72] Boyd was in England at the time, and agreed to attend the recordings. This initial session was followed by further recordings in July. In his 2006 autobiography, the producer recalled being taken aback at Drake's anger and bitterness: "[He said that] I had told him he was a genius, and others had concurred. Why wasn't he famous and rich? This rage must have festered beneath that inexpressive exterior for years."^[73] Both Boyd and Wood noticed a discernible deterioration in Drake's performance, requiring him to overdub his voice separately over the guitar. However, the return to Sound Techniques studio raised Drake's spirits; his mother later recalled, "We were so absolutely thrilled to think that Nick was happy because there hadn't been any happiness in Nick's life for years."^[47]

Death[link]

Drake's gravestone is inscribed with the epitaph 'Now we rise / And we are everywhere', taken from "From the Morning," the final song on his final album, Pink Moon.^[74]

By autumn 1974, Drake's weekly retainer from Island had ceased, and his illness meant he remained in contact with only a few close friends. He had tried to stay in touch with Sophia Ryde, whom he had first met in London in 1968.^[75] Ryde has been described by Drake's biographers as "the nearest thing" to a girlfriend in his life, but she now prefers the description "best (girl) friend".^[76] In a 2005 interview, Ryde revealed that a week before he died, she had sought to end the relationship: "I couldn't cope with it. I asked him for some time. And I never saw him again."^[77] As with the relationship he had earlier shared with fellow folk musician Linda Thompson, Drake's relationship with Ryde was never consummated.^[77]

At some time during the night of 24/25 November 1974, Nick Drake died at home in Far Leys, Tanworth-in-Arden, from an overdose of amitriptyline, a type of antidepressant. He had gone to bed early the night before, after spending the afternoon visiting a friend. His mother stated that, around dawn, he left his room for the kitchen. His family was used to hearing him do this many times before but, during this instance, he did not make a sound. They presumed that he was eating a bowl of cereal. He returned to his room a short while later, and took some pills "to help him sleep".^[78] Drake was accustomed to keeping his own hours; he frequently had difficulty sleeping, and would often stay up through the night playing and listening to music, then sleep late into the following morning. Recalling the events of that night, his mother later stated: "I never used to disturb him at all. But it was about 12 o'clock, and I went in, because really it seemed it was time he got up. And he was lying across the bed. The first thing I saw was his long, long legs."^[79] There was no suicide note, although a letter addressed to Ryde was found close to his bed.^[80]

At the inquest that December, Drake's coroner stated that the cause of death was as a result of "Acute amitriptyline poisoning—self-administered when suffering from a depressive illness", and concluded a verdict of suicide. Though this has been disputed by some members of his family,^[2] there is a general view that accidental or not, Drake had by then given up on life.^[41] Rodney described his son's death as unexpected and extraordinary; however, in a 1979 interview he admitted to "always [being] worried about Nick being so depressed. We used to hide away the aspirin and pills and things like that."^[77] Boyd has stated that he prefers to believe the overdose was accidental. He recalled that Drake's parents had described his mood in the preceding weeks as having been very positive, and that he had planned to move back to London to restart his music career. Boyd believes that this levity was followed by a "crash back into despair". Reasoning that Drake may have taken a high dosage of his antidepressants to recapture this sense of optimism, he said he prefers to imagine Drake "making a desperate lunge for life rather than a calculated surrender to death".^[81] Writing in 1975, NME journalist Nick Kent comments on the irony of Drake's death at a time when he had just begun to regain a sense of "personal balance".^[68] In contrast, Gabrielle Drake has said she prefers to think Drake committed suicide, "in the sense that I'd rather he died because he wanted to end it than it to be the result of a tragic mistake. That would seem to me to be terrible."^[77]

On 2 December 1974, after a service in the Church of St Mary Magdalene, Tanworth-in-Arden, Drake's remains were cremated at the Solihull Crematorium and his ashes later interred under an oak tree in the adjoining graveyard of St Mary's.^[82] The funeral was attended by around 50 mourners, including friends from Marlborough, Aix, Cambridge, London, Witchseason, and Tanworth.^[83] Referring to Drake's tendency to compartmentalise relationships, Brian Wells later observed that many met each other for the first time that morning.^[84] Molly recalled "a lot of his young friends came up here. We'd never met many of them."^[83]

Posthumous popularity[link]

During the early Eighties, I drifted away from the music scene. When I returned, I was surprised to find that Nick Drake was becoming famous.

—Ian MacDonald^[4]

There were no documentaries or compilation albums in the wake of Drake's death.^[85] His public profile remained low throughout the mid and late 1970s, although occasional mentions of his name appeared in the music press. By this time, his parents were receiving an increasing number of fans and admirers as visitors to the family home in Far Leys. Island Records, following a 1975 NME article written by Nick Kent, stated "we have no intention of repackaging Nick's three albums (which remained available), either now or at anytime in the foreseeable future",^[86] but in 1979 Rob Partridge joined Island Records as press officer and commissioned the release of the Fruit Tree box set. Partridge was a fan of Drake's, and had seen him perform early in 1969: "The first thing I did when I got to Island was suggest we put together a retrospective—the studio albums plus whatever else was there. I wasn't necessarily expecting massive vaults with millions of tunes, live recordings or whatever, but there was very little". The release brought together the three studio albums as well as the four tracks recorded with Wood in 1974 and was accompanied by an extensive biography written by the American journalist Arthur Lubow. However, sales were poor and the album received little press notice, and in 1983 Island deleted Fruit Tree from its catalogue.^[14]

By the mid-1980s Drake was being cited as an influence by musicians such as R.E.M.'s Peter Buck and Robert Smith of The Cure. Smith credited the origin of his band's name to a lyric from Drake's song "Time Has Told Me" ("a troubled cure for a troubled mind").^[87] Drake gained further exposure in 1985 with the release of The Dream Academy's hit single "Life in a Northern Town", which included an on-sleeve dedication to Drake.^[88] In 1986 the first biography of Drake was published, in Danish.^[89] His reputation continued to grow, and by the end of the 1980s, his name was appearing regularly in newspapers and music magazines in the United Kingdom;^[90] he had to many come to represent a "doomed romantic hero",^[91] and an "enigma wrapped inside a mystery".^[38] The first step in translating that reputation into record sales came with the release of the compilation album Way to Blue: An Introduction to Nick Drake in May 1994. Although the album never charted in the UK, it sold consistently over the next few years, gaining a gold disc certification in September 1999 for sales of 100,000 copies in the UK.^[92]

On 20 June 1998 BBC Radio 2 broadcast a documentary entitled Fruit Tree: The Nick Drake Story, featuring interviews with Joe Boyd, John Wood, Gabrielle and Molly Drake, Paul Wheeler, Robert Kirby and Ashley Hutchings, and narrated by Danny Thompson.^[93] To tie in with the release of the compilation album Made to Love Magic, an updated version of the documentary was broadcast on 22 May 2004 on Radio 2, retitled Lost Boy: In Search of Nick Drake and featuring the same interview clips but with Thompson's narration replaced by that of Brad Pitt, a self-confessed Nick Drake fan.^[94][95] In early 1999, BBC2 aired a 40-minute documentary, A Stranger Among Us—In Search of Nick Drake. The following year, Dutch director Jeroen Berkvens released the documentary A Skin Too Few: The Days of Nick Drake, featuring interviews with Boyd, Gabrielle Drake, Wood and Kirby. Later that year, The Guardian placed Bryter Layter at number 1 in its "Alternative top 100 albums ever" list.^[70]

In 1999, "Pink Moon" was used in "Milky Way", a Volkswagen Cabrio commercial, leading to a large increase in record sales.^[96][97] The US telecommunications company AT&T used "From the Morning" in one of their advertisements as part of their "Rethink Possible" campaign in North America in 2010.^[96]

In recent years several musicians, including Lucinda Williams, Badly Drawn Boy, Lou Barlow and Mikael Åkerfeldt have cited Drake as an influence. In 2004, nearly 30 years after his death, Drake gained his first chart placing when two singles, "Magic" and "River Man", were released to coincide with the Made to Love Magic album.

On 16 May 2009 Joe Boyd curated a concert at the Birmingham Town Hall, with Robert Kirby as musical arranger and a variety of singers and musicians performing Drake's songs. Among the featured artists were Robyn Hitchcock, Camille O'Sullivan, Martha Wainwright, Graham Coxon, Beth Orton, Harper Simon, Kate St John, Stuart Murdoch and Vashti Bunyan.^[98] Following this concert's success, Boyd staged a short concert tour of the UK in January 2010, following the same format: artists on the tour included Vashti Bunyan, Green Gartside, Lisa Hannigan, Scott Matthews, Teddy Thompson, Krystle Warren, Robyn Hitchcock, Kirsty Almeida and Harper Simon. The concert at the London Barbican Centre was filmed and broadcast on BBC Four in April 2010.^[99]

Musical and lyrical style[link]

Drake was obsessive about practicing his guitar technique, and would often stay up through the night experimenting with tunings and working on songs. His mother remembered hearing him "bumping around at all hours. I think he wrote his nicest melodies in the early-morning hours."^[14] Self-taught,^[48] he achieved his guitar style through the use of alternative tunings to create cluster chords.^[100] These are difficult to achieve on a guitar using standard tuning; Drake used tunings which made cluster chords available using more conventional chord shapes. In many songs he accents the dissonant effect of such non-standard tunings through his vocal melodies.^[100]

Drake studied English literature in Cambridge and was particularly drawn to the works of William Blake, William Butler Yeats and Henry Vaughan, and his lyrics reflect such influences.^[4] Drake also employs a series of elemental^[101] symbols and codes, largely drawn from nature. The moon, stars, sea, rain, trees, sky, mist and seasons are all commonly used, influenced in part by his rural upbringing.^[4] Images related to summer figure centrally in his early work; from Bryter Layter on, his language is more autumnal, evoking a season commonly used to convey senses of loss and sorrow.^[4] Throughout, Drake writes with detachment, more as an observer than participant, a point of view Rolling Stone's Anthony DeCurtis described "as if he were viewing his life from a great, unbridgeable distance."^[101] This perceived inability to connect has led to much speculation about Drake's sexuality.^[102] Boyd has said he detects a virginal quality in his lyrics and music, and notes that he never observed or heard of the singer behaving in a sexual way with anyone, male or female.^[103] Kirby described Drake's lyrics as a "series of extremely vivid, complete observations, almost like a series of epigrammatic proverbs", though he doubts that Drake saw himself as "any sort of poet". Instead he believes that Drake's lyrics were crafted to "complement and compound a mood that the melody dictates in the first place."^[68]

Discography[link]

References[link]

Notes[link]