A deer and two fawns feeding on foliage

Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs^{[1][page needed]} such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in general are known as primary consumers.

Herbivory usually refers to animals eating plants; fungi, bacteria and protists that feed on living plants are usually termed plant pathogens (plant diseases),and microbes that feed on dead plants are saprotrophs. Flowering plants that obtain nutrition from other living plants are usually termed parasitic plants.

A herbivore is not the same as a vegetarian, a human who voluntarily undertakes a primarily herbivorous diet.^[2][3]

Etymology[link]

Herbivore is the anglicized form of a modern Latin coinage, herbivora, cited in Charles Lyell's 1830 Principles of Geology.^[4] Richard Owen employed the anglicized term in an 1854 work on fossil teeth and skeletons.^[4] Herbivora is derived from the Latin herba meaning a small plant or herb,^[5] and vora, from vorare, to eat or devour.^[6]

[edit] Evolution of herbivory

A fossil Viburnum lesquereuxii leaf with evidence of insect herbivory; Dakota Sandstone (Cretaceous) of Ellsworth County, Kansas. Scale bar is 10 mm.

Our understanding of herbivory in geological time comes from three sources: fossilized plants, which may preserve evidence of defence (such as spines), or herbivory-related damage; the observation of plant debris in fossilised animal faeces; and the construction of herbivore mouthparts.^[7]

Long thought to be a Mesozoic phenomenon, evidence for herbivory is found almost as soon as fossils which could show it. Within under 20 million years of the first land plants evolving, plants were being consumed by insects.^[8] Insects fed on the spores of early Devonian plants, and the Rhynie chert also provides evidence that organisms fed on plants using a "pierce and suck" technique.^[7]

Herbivory among terrestrial vertebrates (tetrapods) came much later. Early tetrapods were large amphibious piscivores. While amphibians continued to feed on fish and later insects, reptiles began exploring two new food types, tetrapods (carnivory), and later, plants (herbivory). Carnivory was a natural transition from insectivory for medium and large tetrapods, requiring minimal adaptation. In contrast, a complex set of adaptations was necessary for feeding on highly fibrous plant materials).^[9]

During the ensuing 75 million years^{[citation needed]}, plants evolved a range of more complex organs - from roots to seeds. There is no evidence for these being fed upon until the middle-late Mississippian, 326.4 million years ago. There was a gap of 50 to 100 million years between each organ evolving, and it being fed upon; this may be due to the low levels of oxygen during this period, which may have suppressed evolution.^[8] Further than their arthropod status, the identity of these early herbivores is uncertain.^[8] Hole feeding and skeletonisation are recorded in the early Permian, with surface fluid feeding evolving by the end of that period.^[7]

Arthropods have evolved herbivory in four phases, changing their approach to herbivory in response to changing plant communities.^[10]
Another stage of herbivore evolution is characterized by the evolution of tetrapod herbivores, with the first appearance in the fossil record near the Permio-Carboniferous boundary approximately 300 MYA. The earliest evidence of herbivory by tetrapod organisms is seen in fossils of jawbones where dental occlusion (process by which teeth from the upper jaw come in contact with those in the lower jaw) is present. The evolution of dental occlusion lead to a drastic increase in food processing associated with herbivory and provides direct evidence about feeding strategies based on tooth wear patterns. Examination of phylogenetic frameworks reveals that dental occlusion developed independently in several lineages through dental and mandibular morphologes, suggesting that the evolution and radiation of tetrapod herbivores occurred simultaneously within various lineages.^[11]

Food chain[link]

Leaf miners feed on leaf tissue between the epidermal layers, leaving visible trails

Herbivores form an important link in the food chain as they consume plants in order to receive the carbohydrates produced by a plant from photosynthesis. Carnivores in turn consume herbivores for the same reason, while omnivores can obtain their nutrients from either plants or animals. Due to a herbivore's ability to survive solely on tough and fibrous plant matter, they are termed the primary consumers in the food cycle(chain). Herbivory, carnivory, and omnivory can be regarded as special cases of Consumer-Resource Systems.^[12]

Predator-prey theory (herbivore-plant interactions)[link]

According to the theory of predator-prey interactions, the relationship between herbivores and plants is cyclic.^[13] When prey (plants) are numerous their predators (herbivores) increase in numbers, reducing the prey population, which in turn causes predator number to decline.^[14] The prey population eventually recovers, starting a new cycle. This suggests that the population of the herbivore fluctuates around the carrying capacity of the food source, in this case the plant.

Several factors play into these fluctuating populations and help stabilize predator-prey dynamics. For example, spatial heterogeneity is maintained, which means there will always be pockets of plants not found by herbivores. This stabilizing dynamic plays an especially important role for specialist herbivores that feed on one species of plant and prevents these specialists from wiping out their food source.^[15] Prey defenses also help stabilize predator-prey dynamic, and for more information on these relationships see the section on Plant Defenses. Eating a second prey type helps herbivores’ populations stabilize.^[16] Alternating between two or more plant types provides population stability for the herbivore, while the populations of the plants oscillate.^[17] This plays an important role for generalist herbivores that eat variety of plants. Keystone herbivores keep vegetation populations in check and allow for a greater diversity of both herbivores and plants.^[16] When an invasive herbivore or plant enters the system, the balance is thrown off and the diversity can collapse to a monotaxon system.^[16]

Feeding strategies[link]

Herbivores are limited in their feeding ability by either time or resources. Animals that are time limited, meaning they have a limited amount of time to consume the food they need, use a feeding strategy of grazing and browsing, while those animals that are resource limited, meaning that they are limited in the type of food they eat, use a selective feeding strategy. Grazers/browsers tend to be either very large herbivores that need to consume a lot of food in order to maintain their metabolism, or herbivores that have a very short amount of time to eat as much as possible before reproducing, like many generalist insects. Several theories attempt to explain and quantify the relationship between animals and their food, such as Kleiber's law, Holling's disk equation and Marginal Value Theorem.

Kleiber's law explains the relationship between the size of the animal and the feeding strategy it uses. In essence, it says that larger animals need to eat less food, per unit weight, than smaller animals.^[18] Kleiber’s law states that the metabolic rate (q₀) of an animal is the mass of the animal (M) raise to the 3/4 power:
q₀=M^3/4
Therefore, the mass of the animal increases at a faster rate than the metabolic rate.^[19]
There are many types of feeding strategies employed by herbivores. Many herbivores do not fall into one specific feeding strategy, but instead employ several strategies and eat a variety of plant parts.

Types of feeding strategies:

Optimal Foraging Theory is a model for predicting animal behavior while looking for food or other resource, such as shelter or water. This model assesses both individual movement, such as animal behavior while looking for food, and distribution within a habitat, such as dynamics at the population and community level. For example, the model would be used to look at the browsing behavior of a deer while looking for food, as well as that deer's specific location and movement within the forested habitat and its interaction with other deer while in that habitat.

This model can be controversial, where critics say that the theory is circular and untestable. Critics say that the theory uses examples that fit the theory, but that researchers do not use the theory when it does not fit the reality.^[20][21] Other critics point out that animals do not have the ability to assess and maximize their potential gains, therefore the optimal foraging theory is irrelevant and derived to explain trends that do not exist in nature.^[22][23]

Holling's disk equation models the efficiency at which predators consume prey. The model predicts that as the number of prey increases, the amount of time predators spend handling prey also increases and therefore the efficiency of the predator decreases.^[24] In 1959, S. Holling proposed an equation to model the rate of return for an optimal diet: Rate (R ) = Energy gained in foraging (Ef)/(time searching (Ts) + time handling (Th))
Failed to parse (Missing texvc executable; please see math/README to configure.): R = Ef/(Ts + Th)
Where s = cost of search per unit time f = rate of encounter with items, h = handling time, e = energy gained per encounter
In effect, this would indicate that a herbivore in a dense forest would spend more time getting handling (eating) the vegetation because there was so much vegetation around than a herbivore in a sparse forest, who could easily browse through the forest vegetation. Therefore, according to the Holling's disk equation, the herbivore in the sparse forest would be more efficient at eating than the herbivore in the dense forest

Marginal Value Theorem describes the balance between eating all the food in a patch for immediate energy, or moving to a new patch and leaving the plants in the first patch to regenerate for future use. The theory predicts that absent complicating factors, an animal should leave a resource patch when the rate of payoff (amount of food) falls below the average rate of payoff for the entire area.^[25] According to this theory, therefore, locus should move to a new patch of food when the patch they are currently feeding on requires more energy to obtain food than an average patch. Within this theory, two subsequent parameters emerge, the Giving Up Density (GUD) and the Giving Up Time (GUT). The Giving Up Density (GUD) quantifies the amount of food that remains in a patch when a forager moves to a new patch.^[26] The Giving Up Time (GUT) is used when an animal continuously assesses the patch quality.^[27]

Attacks and counter-attacks[link]

Plant defense[link]

A plant defense is a trait that increases plant fitness when faced with herbivory. This is measured relative to another plant that lacks the defensive trait. Plant defenses increase survival and/or reproduction (fitness) of plants under pressure of predation from herbivores.

Defense can be divided into two main categories, tolerance and resistance. Tolerance is the ability of a plant to withstand damage without a reduction in fitness. This can occur by diverting herbivory to non-essential plant parts or by rapid regrowth and recovery from herbivory. Resistance refers to the ability of a plant to reduce the amount of damage it receives from a herbivore. This can occur via avoidance in space or time,^[28] physical defenses, or chemical defenses. Defenses can either be constitutive, always present in the plant, or induced, produced or translocated by the plant following damage or stress.^[29]

Physical, or mechanical, defenses are barriers or structures designed to deter herbivores or reduce intake rates, lowering overall herbivory. thorns such as those found on roses or acacia trees are one example, as are the spines on a cactus. Smaller hairs known as trichomes may cover leaves or stems and are especially effective against invertebrate herbivores.^[30] In addition, some plants have waxes or resins that alter their texture, making them difficult to eat. Finally, some plants sequester silica inside their tissues. These are analogous to small pieces of glass that wear down the teeth of herbivores^{[citation needed]}.

Chemical defenses are secondary metabolites produced by the plant that deter herbivory. There are a wide variety of these in nature and a single plant can have hundreds of different chemical defenses. Chemical defenses can be divided into two main groups, carbon-based defenses and nitrogen-based defenses.

Carbon-based defenses include terpenes and phenolics. Terpenes are derived from 5-carbon isoprene units and comprise essential oils, carotenoids, resins, and latex. They can have a number of functions that disrupt herbivores such as inhibiting adenosine triphosphate (ATP) formation, molting hormones, or the nervous system.^[31] Phenolics combine an aromatic carbon ring with a hydroxyl group. There are a number of different phenolics such as lignins, which are found in cell walls and are very indigestible except for specialized microorgamisms; tannins, which have a bitter taste and bind to proteins making them indigestible; and furanocumerins, which produce free radicals disrupting DNA, protein, and lipids, and can cause skin irritation.

Nitrogen-based defenses are synthesized from amino acids and primarily come in the form of alkaloids and cyanogens. Alkaloids include commonly recognized substances such as caffeine, nicotine, and morphine. These compounds are often bitter and can inhibit DNA or RNA synthesis or block nervous system signal transmission. Cyanogens get their name from the cyanide stored within their tissues. This is released when the plant is damaged and inhibits cellular respiration and electron transport.

Plants have also changed features that enhance the probability of attracting natural enemies to herbivores. Some emit semiochemicals, odors that attract natural enemies, while others provide food and housing to maintain the natural enemies’ presence (e.g. ants that reduce herbivory^[32]). A given plant species often has many types of defensive mechanisms, mechanical or chemical, constitutive or induced, which additively serve to protect the plant, and allow it to escape from herbivores.

Herbivore offense[link]

Aphids are fluid feeders on plant sap.

The myriad of defenses displayed by plants means that their herbivores need a variety of techniques to overcome these defenses and obtain food. These allow herbivores to increase their feeding and use of a host plant. Herbivores have three primary strategies for dealing with plant defenses: choice, herbivore modification, and plant modification.

Feeding choice involves which plants a herbivore chooses to consume. It has been suggested that many herbivores feed on a variety of plants to balance their nutrient uptake and to avoid consuming too much of any one type of defensive chemical. This involves a tradeoff however, between foraging on many plant species to avoid toxins or specializing on one type of plant that can be detoxified.^[33]

Herbivore modification is when various adaptations to body or digestive systems of the herbivore allow them to overcome plant defenses. This might include detoxifying secondary metabolites,^[34] sequestering toxins unaltered,^[35] or avoiding toxins, such as through the production of large amounts of saliva to reduce effectiveness of defenses. Herbivores may also utilize symbionts to evade plant defences. For example, some aphids use bacteria in their gut to provide essential amino acids lacking in their sap diet.^[36]

Plant modification occurs when herbivores manipulate their plant prey to increase feeding. For example, some caterpillars roll leaves to reduce the effectiveness of plant defenses activated by sunlight.^[37]

The adaptation dance[link]

The back and forth relationship of plant defense and herbivore offense can be seen as a sort of "adaptation dance" in which one partner makes a move and the other counters it.^[34] This reciprocal change drives coevolution between many plants and herbivores, resulting in what has been referred to as a "coevolutionary arms race".^[38] The escape and radiation mechanisms for coevolution, presents the idea that adaptations in herbivores and their host plants, has been the driving force behind speciation.^[39][40]

It is important to remember that while much of the interaction of herbivory and plant defense is negative, with one individual reducing the fitness of the other, some is actually beneficial. This beneficial herbivory takes the form of mutualisms in which both partners benefit in some way from the interaction. Seed dispersal by herbivores and pollination are two forms of mutualistic herbivory in which the herbivore receives a food resource and the plant is aided in reproduction.^[41]

Impacts of herbivores[link]

The impact of herbivory can be seen in many areas ranging from economics to ecological, and sometimes affecting both. For example, environmental degradation from white-tailed deer (Odocoileus virginianus) in the U.S. alone has the potential to both change vegetative communities through over-browsing and cost forest restoration projects upwards of $750 million annually. Agricultural crop damage by the same species totals approximately $100 million every year. Insect crop damages also contribute largely to annual crop losses in the U.S.^[42] Another area in which herbivory greatly affects economics is through the revenue generated by recreational uses of herbivorous organisms, such as hunting and ecotourism. For example, the hunting of herbivorous game species such as white-tailed deer, cottontail rabbits, antelope, and elk in the U.S. contributes greatly to the billion-dollar annually hunting industry. Ecotourism is another major source of revenue, particularly in Africa, where many large mammalian herbivores such as elephants, zebras, and giraffes help to bring in the equivalent of millions of US dollars to various nations annually.

See also[link]

Wikimedia Commons has media related to: Herbivory

• Browsing (herbivory)
• Carnivore
• Consumer-resource systems
• Grazing
• List of feeding behaviours
• List of herbivorous animals
• Omnivore
• Plant-based diet (disambiguation)
• Pollination
• Productivity (ecology)
• Seed dispersal
• Seed predation

References[link]

Further reading[link]

• Bob Strauss, 2008, Herbivorous Dinosaurs, The New York Times
• Danell, K., R. Bergström, P. Duncan, J. Pastor (Editors)(2006) Large herbivore ecology, ecosystem dynamics and conservation Cambridge, UK : Cambridge University Press. 506 p. ISBN 0-521-83005-2
• Crawley, M. J. (1983) Herbivory : the dynamics of animal-plant interactions Oxford : Blackwell Scientific. 437 p. ISBN 0-632-00808-3
• Olff, H., V.K. Brown, R.H. Drent (editors) (1999) Herbivores : between plants and predators Oxford ; Malden, Ma. : Blackwell Science. 639 p. ISBN 0-632-05155-8

External links[link]

• Herbivore information resource website
• The herbivore defenses of Senecio viscusus
• Herbivore defense in Lindera benzoin
• website of the herbivory lab at Cornell University

v t e Feeding behaviours Carnivores adult Hematophagy Insectivore Lepidophagy Man-eater Molluscivore Mucophagy Myrmecophagy Ophiophagy Piscivore Avivore Spongivore Vermivore Herpetivore reproductive Oophagy Paedophagy Placentophagy Breastfeeding Weaning cannibalistic Cannibalism Human cannibalism Self-cannibalism Sexual cannibalism Herbivores Folivore Frugivore Graminivore Granivore Nectarivore Palynivore Xylophagy Osteophagy Cellular Phagocytosis Myzocytosis Others Bacterivore Fungivore Coprophagia Detritivore Geophagy Omnivore Methods Apex predator Bait balls Bottom feeding Browsing Feeding frenzy Filter feeding Grazing Hypercarnivore Intraguild predation Kleptoparasitism Scavenging Trophallaxis Predation Antipredator adaptation Carnivorous plant Carnivorous fungus Carnivorous protist Category:Eating behaviors

v t e Modelling ecosystems – trophic components General Abiotic component Abiotic stress Behaviour Biogeochemical cycle Biomass Biotic component Biotic stress Carrying capacity Competition Ecosystem Ecosystem ecology Ecosystem model Keystone species List of feeding behaviours Metabolic theory of ecology Productivity Producers Autotrophs Chemosynthesis Chemotrophs Foundation species Mixotrophs Myco-heterotrophy Mycotroph Organotrophs Photoheterotrophs Photosynthesis Photosynthetic efficiency Phototrophs Primary nutritional groups Primary production Consumers Apex predator Bacterivore Carnivores Chemoorganotroph Foraging Generalist and specialist species Intraguild predation Herbivores Heterotroph Heterotrophic nutrition Insectivore Mesopredator release hypothesis Omnivores Optimal foraging theory Predation Prey switching Decomposers Chemoorganoheterotrophy Decomposition Detritivores Detritus Microorganisms Archaea Bacteriophage Environmental microbiology Lithoautotroph Lithotrophy Microbial cooperation Microbial ecology Microbial food web Microbial intelligence Microbial loop Microbial mat Microbial metabolism Phage ecology Food webs Biomagnification Ecological efficiency Ecological pyramid Energy flow Food chain Trophic level Example webs Cold seeps Hydrothermal vents Intertidal Kelp forests Lakes North Pacific Subtropical Gyre Rivers San Francisco Estuary Soil Tidal pool Processes Ascendency Bioaccumulation Cascade effect Climax community Competitive exclusion principle Consumer-resource systems Copiotrophs Dominance Ecological network Ecological succession Energy quality Energy Systems Language f-ratio Feed conversion ratio Feeding frenzy Mesotrophic soil Nutrient cycle Oligotroph Paradox of the plankton Trophic cascade Trophic mutualism Trophic state index Defense/counter Animal coloration Antipredator adaptations Camouflage Herbivore adaptations to plant defense Mimicry Plant defense against herbivory Predator avoidance in schooling fish

v t e Modelling ecosystems – other components Population ecology Abundance Allee effect Depensation Ecological yield Effective population size Intraspecific competition Logistic function Malthusian growth model Maximum sustainable yield Overpopulation in wild animals Overexploitation Population cycle Population dynamics Population modeling Population size Predator–prey equations Recruitment Resilience Small population size Stability Species Biodiversity Density-dependent inhibition Ecological effects of biodiversity Ecological extinction Endemic species Flagship species Gradient analysis Indicator species Introduced species Invasive species Latitudinal gradients in species diversity Minimum viable population Neutral theory Occupancy-abundance relationship Population viability analysis Priority effect Rapoport's rule Relative abundance distribution Relative species abundance Species diversity Species homogeneity Species richness Species distribution Species-area curve Umbrella species Species interaction Antibiosis Biological interaction Commensalism Community ecology Ecological facilitation Interspecific competition Mutualism Storage effect Spatial ecology Biogeography Cross-boundary subsidy Ecocline Ecotone Ecotype Disturbance Edge effect Foster's rule Habitat fragmentation Ideal free distribution Intermediate Disturbance Hypothesis Island biogeography Landscape ecology Landscape epidemiology Landscape limnology Metapopulation Patch dynamics r/K selection theory Source–sink dynamics Niche Ecological niche Ecological trap Ecosystem engineer Environmental niche modelling Guild Habitat Limiting similarity Niche apportionment models Niche construction Niche differentiation Other networks Assembly rules Bateman's principle Bioluminescence Ecological collapse Ecological debt Ecological deficit Ecological energetics Ecological indicator Ecological threshold Ecosystem diversity Emergence Extinction debt Kleiber's law Liebig's law of the minimum Marginal value theorem Thorson's rule Xerosere Other Allometry Alternative stable state Balance of nature Biological data visualization Constructal theory Ecocline Ecological economics Ecological footprint Ecological forecasting Ecological humanities Ecological stoichiometry Ecopath Ecosystem based fisheries Endolith Evolutionary ecology Functional ecology Industrial ecology Macroecology Microecosystem Natural environment Systems ecology Theoretical ecology List of ecology topics

Sir David Attenborough
Attenborough in May 2003
Born	(1926-05-08) 8 May 1926 (age 86) Isleworth, London
Nationality	British
Alma mater	Clare College, Cambridge (natural sciences) London School of Economics (social anthropology)
Occupation	Broadcaster Naturalist
Title	Member of the Order of Merit Companion of Honour Commander of the Royal Victorian Order Commander of the Order of the British Empire Fellow of the Royal Society Fellow of Zoological Society of London
Spouse	Jane Elizabeth Ebsworth Oriel (m. 1950–1997, her death)
Children	Robert Attenborough Susan Attenborough

Sir David Frederick Attenborough ( /ˈætənbərə/) OM, CH, CVO, CBE, FRS, FZS, FSA (born 8 May 1926) is a British broadcaster and naturalist.

His career as the face and voice of natural history programmes has endured for more than 50 years. He is best known for writing and presenting the nine Life series, in conjunction with the BBC Natural History Unit, which collectively form a comprehensive survey of all life on the planet. He is also a former senior manager at the BBC, having served as controller of BBC Two and director of programming for BBC Television in the 1960s and 1970s.

Attenborough is widely considered a national treasure in Britain, although he himself does not care for the term.^[1][2][3] He is a younger brother of director, producer and actor Richard Attenborough.^[4]

Early life and family[link]

Attenborough was born in Isleworth, west London, but grew up in College House on the campus of the University College, Leicester, where his father, Frederick, was principal.^[5] He is the middle of three sons (his elder brother, Richard, became an actor and his younger brother, John, an executive at Italian car manufacturer Alfa Romeo).^[6] During World War II his parents also adopted two Jewish refugee girls from Europe.^[7][8]

Attenborough spent his childhood collecting fossils, stones and other natural specimens. He received encouragement in this pursuit at age seven, when a young Jacquetta Hawkes admired his "museum". A few years later, one of his adoptive sisters gave him a piece of amber filled with prehistoric creatures; some 50 years later, it would be the focus of his programme The Amber Time Machine.

Attenborough was educated at Wyggeston Grammar School for Boys in Leicester and then won a scholarship to Clare College, Cambridge in 1945, where he studied geology and zoology and obtained a degree in natural sciences.^[9] In 1947, he was called up for national service in the Royal Navy and spent two years stationed in North Wales and the Firth of Forth.

In 1950, Attenborough married Jane Elizabeth Ebsworth Oriel; the marriage lasted until her death in 1997. The couple had two children, Robert and Susan.^[8] Robert is a senior lecturer in bioanthropology for the School of Archaeology and Anthropology at the Australian National University in Canberra.^[10]

First years at the BBC[link]

After leaving the Navy, Attenborough took a position editing children's science textbooks for a publishing company. He soon became disillusioned with the work, however, and in 1950, he applied for a job as a radio talks producer with the BBC. Although he was rejected for this job, his CV later attracted the interest of Mary Adams, head of the Talks (factual broadcasting) department of the BBC's fledgling television service. Attenborough, like most Britons at that time, did not own a television, and he had seen only one programme in his life.^[11] However, he accepted Adams' offer of a three-month training course, and in 1952 he joined the BBC full-time. Initially discouraged from appearing on camera because Adams thought his teeth were too big,^[12] he became a producer for the Talks department, which handled all non-fiction broadcasts. His early projects included the quiz show Animal, Vegetable, Mineral? and Song Hunter, a series about folk music presented by Alan Lomax.

Attenborough's association with natural history programmes began when he produced and presented the three-part series The Pattern of Animals. The studio-bound programme featured animals from London Zoo, with the naturalist Julian Huxley discussing their use of camouflage, aposematism and courtship displays. Through this programme, Attenborough met Jack Lester, the curator of the zoo's reptile house, and they decided to make a series about an animal-collecting expedition. The result was Zoo Quest, first broadcast in 1954, which Attenborough presented at short notice, due to Lester being taken ill.

In 1957, the BBC Natural History Unit was formally established in Bristol. Attenborough was asked to join it, but declined, not wishing to move from London where he and his young family were settled. Instead he formed his own department, the Travel and Exploration Unit,^[13] which allowed him to continue to front Zoo Quest as well as produce other documentaries, notably the Travellers' Tales and Adventure series.

In the early 1960s, Attenborough resigned from the permanent staff of the BBC to study for a postgraduate degree in social anthropology at the London School of Economics, interweaving his study with further filming.^[14] However, he accepted an invitation to return to the BBC as controller of BBC Two before he could finish the degree.

BBC administration[link]

Attenborough became the controller of BBC Two in March 1965, but had a clause inserted in his contract that would allow him to continue making programmes on an occasional basis. Later the same year, he filmed elephants in Tanzania, and in 1969, he made a three-part series on the cultural history of the Indonesian island of Bali. For the 1971 film A Blank on the Map, he joined the first Western expedition to a remote highland valley in New Guinea to seek out a lost tribe.

BBC Two was launched in 1964, but had struggled to capture the public's imagination. When Attenborough arrived as controller, he quickly abolished the channel's quirky kangaroo mascot and shook up the schedule. With a mission to make BBC Two's output diverse and different from that offered by other networks, he began to establish a portfolio of programmes that defined the channel's identity for decades to come. Under his tenure, music, the arts, entertainment, archaeology, experimental comedy, travel, drama, sport, business, science and natural history all found a place in the weekly schedules. Often, an eclectic mix was offered within a single evening's viewing. Programmes he commissioned included Man Alive, Call My Bluff, Chronicle, Life, One Pair of Eyes, The Old Grey Whistle Test, Monty Python's Flying Circus and The Money Programme. When BBC Two became the first British channel to broadcast in colour in 1967, Attenborough took advantage by introducing televised snooker, as well as bringing rugby league to British television on a regular basis via the BBC2 Floodlit Trophy.

One of his most significant decisions was to order a 13-part series on the history of Western art, to show off the quality of the new UHF colour television service that BBC Two offered. Broadcast to universal acclaim in 1969, Civilisation set the blueprint for landmark authored documentaries, which were informally known as "tombstone" or "sledgehammer" projects. Others followed, including Jacob Bronowski's The Ascent of Man (also commissioned by Attenborough), and Alistair Cooke's America. Attenborough thought that the story of evolution would be a natural subject for such a series. He shared his idea with Chris Parsons, a producer at the Natural History Unit, who came up with the title Life on Earth and returned to Bristol to start planning the series. Attenborough harboured a strong desire to present the series himself, but this would not be possible so long as he remained in a management post.

In 1969, Attenborough was promoted to director of programmes, making him responsible for the output of both BBC channels. His tasks, which included agreeing budgets, attending board meetings and firing staff, were now far removed from the business of filming programmes. When Attenborough's name was being suggested as a candidate for the position of Director General of the BBC in 1972, he phoned his brother Richard to confess that he had no appetite for the job. Early the following year, he left his post to return to full-time programme-making, leaving him free to write and present the planned natural history epic.

Return to broadcasting[link]

After his resignation, Attenborough became a freelance broadcaster and immediately started work on his next project, a pre-arranged trip to Indonesia with a crew from the Natural History Unit. It resulted in the 1973 series Eastwards with Attenborough, which was similar in tone to the earlier Zoo Quests but without the animal-collecting element.

On his return, he began to work on the scripts for Life on Earth. Due to the scale of his ambition, the BBC decided to partner with an American network to secure the necessary funding. While the negotiations were proceeding he worked on a number of other television projects. He presented a series on tribal art (The Tribal Eye, 1975) and another on the voyages of discovery (The Explorers, 1975). He also presented a BBC children's series about cryptozoology entitled Fabulous Animals (1975), which featured mythical creatures such as the griffin and kraken.^[15] Eventually, the BBC signed a co-production deal with Turner Broadcasting and Life on Earth moved into production in 1976.

Life series[link]

Beginning with Life on Earth in 1979, Attenborough set about creating a body of work which became a benchmark of quality in wildlife film-making and influenced a generation of documentary film-makers. The series also established many of the hallmarks of the BBC's natural history output. By treating his subject seriously and researching the latest discoveries, Attenborough and his production team gained the trust of the scientific community, who responded by allowing him to feature their subjects in his programmes. In Rwanda, for example, Attenborough and his crew were granted privileged access to film Dian Fossey's research group of mountain gorillas. Innovation was another factor in Life on Earth's success: new film-making techniques were devised to get the shots Attenborough wanted, with a focus on events and animals that were hitherto unfilmed. Computerised airline schedules, which had only recently been introduced, enabled the series to be elaborately devised so that Attenborough visited several locations around the globe in each episode, sometimes even changing continents mid-sentence. Although appearing as the on-screen presenter, he consciously restricted his pieces to camera to give his subjects top billing.

The success of Life on Earth prompted the BBC to consider a follow-up, and five years later, The Living Planet was screened. This time, Attenborough built his series around the theme of ecology, the adaptations of living things to their environment. It was another critical and commercial success, generating huge international sales for the BBC. In 1990, The Trials of Life completed the original Life trilogy, looking at animal behaviour through the different stages of life. The series drew strong reactions from the viewing public for its sequences of killer whales hunting sea lions on a Patagonian beach and chimpanzees hunting and violently killing a colobus monkey.

In the 1990s, Attenborough continued to use the "Life" moniker for a succession of authored documentaries. In 1993, he presented Life in the Freezer, the first television series to survey the natural history of Antarctica. Although past normal retirement age, he then embarked on a number of more specialised surveys of the natural world, beginning with plants. They proved a difficult subject for his producers, who had to deliver five hours of television featuring what are essentially immobile objects. The result, The Private Life of Plants (1995), showed plants as dynamic organisms by using time-lapse photography to speed up their growth.

Prompted by an enthusiastic ornithologist at the BBC Natural History Unit, Attenborough then turned his attention to the animal kingdom and in particular, birds. As he was neither an obsessive twitcher, nor a bird expert, he decided he was better qualified to make The Life of Birds (1998) on the theme of behaviour. The order of the remaining "Life" series was dictated by developments in camera technology. For The Life of Mammals (2002), low-light and infrared cameras were deployed to reveal the behaviour of nocturnal mammals. The series contains a number of memorable two shots of Attenborough and his subjects, which included chimpanzees, a blue whale and a grizzly bear. Advances in macro photography made it possible to capture natural behaviour of very small creatures for the first time, and in 2005, Life in the Undergrowth introduced audiences to the world of invertebrates.

At this point, Attenborough realised that he had spent 20 years unconsciously assembling a collection of programmes on all the major groups of terrestrial animals and plants — only reptiles and amphibians were missing. When Life in Cold Blood was broadcast in 2008, he had the satisfaction of completing the set, brought together in a DVD encyclopaedia called Life on Land. In an interview that year, Attenborough was asked to sum up his achievement, and responded:

The evolutionary history is finished. The endeavour is complete. If you'd asked me 20 years ago whether we'd be attempting such a mammoth task, I'd have said "Don't be ridiculous!" These programmes tell a particular story and I'm sure others will come along and tell it much better than I did, but I do hope that if people watch it in 50 years' time, it will still have something to say about the world we live in.^[16]

However, in 2010 Attenborough asserted that his First Life — dealing with evolutionary history before Life on Earth — should also be included within the "Life" series. In the documentary Attenborough's Journey he stated, "This series, to a degree which I really didn't fully appreciate until I started working on it, really completes the set."^[17]

Other documentaries[link]

Alongside the "Life" series, Attenborough has continued to work on other television documentaries, mainly in the natural history genre. He wrote and presented a series on man's influence on the natural history of the Mediterranean basin, The First Eden, in 1987. Two years later, he demonstrated his passion for fossils in Lost Worlds Vanished Lives.

Attenborough narrated every episode of Wildlife on One, a BBC One wildlife series which ran for 253 episodes between 1977 and 2005. At its peak, it drew a weekly audience of eight to ten million, and the 1987 episode "Meerkats United" was voted the best wildlife documentary of all time by BBC viewers.^[18] He has also narrated over 50 episodes of Natural World, BBC Two's flagship wildlife series. (Its forerunner, The World About Us, was created by Attenborough in 1969, as a vehicle for colour television.^[19]) In 1997, he narrated the BBC Wildlife Specials, each focussing on a charismatic species, and screened to mark the Natural History Unit's 40th anniversary.

As a writer and narrator, he continued to collaborate with the BBC Natural History Unit in the new millennium. Alastair Fothergill, a senior producer with whom Attenborough had worked on The Trials of Life and Life in the Freezer, was making The Blue Planet (2001), the Unit's first comprehensive series on marine life. He decided not to use an on-screen presenter due to difficulties in speaking to camera through diving apparatus, but asked Attenborough to narrate the films. The same team reunited for Planet Earth (2006), the biggest nature documentary ever made for television and the first BBC wildlife series to be shot in high definition. In 2011, Fothergill gave Attenborough a more prominent role in Frozen Planet, a major series on the natural history of the polar regions. Attenborough appeared on screen and authored the final episode, in addition to performing voiceover duties.

In 2009, he co-wrote and narrated Life, a ten-part series focussing on extraordinary animal behaviour,^[20] and narrated Nature's Great Events, which showed how seasonal changes trigger major natural spectacles.^[21]

By the turn of the millennium, Attenborough's authored documentaries were adopting a more overtly environmentalist stance. In State of the Planet (2000), he used the latest scientific evidence and interviews with leading scientists and conservationists to assess the impact of man's activities on the natural world. He later turned to the issues of global warming (The Truth about Climate Change, 2006) and human population growth (How Many People Can Live on Planet Earth?, 2009). He also contributed a programme which highlighted the plight of endangered species to the BBC's Saving Planet Earth project in 2007, the 50th anniversary of the Natural History Unit.

Attenborough is also forging a new partnership with Sky, working on documentaries for the broadcaster's new 3D network, Sky 3D. Their first collaboration was Flying Monsters 3D, a film about pterosaurs which debuted on Christmas Day of 2010.^[22] A second film, The Bachelor King 3D, followed a year later, and further collaborations are planned.

Current projects[link]

Attenborough celebrates his 60th year in broadcasting in 2012, and continues to work on a number of television, film and radio projects. The anniversary will be marked by the BBC with the screening of Attenborough: 60 Years in the Wild, a three part series which looks at how wildlife filmmaking techniques, discoveries in the field of biological sciences and our relationship with the natural world have all developed over the past 60 years.^[23]

Further collaborations with Sky 3D are also in the pipeline. First will come Kingdom of Plants 3D, a three-part series filmed at the Royal Botanical Gardens, Kew which debuts on 26 May 2012 and will be simulcast on Sky Atlantic HD.^[24] A series on the wildlife of the Galapagos Islands has also been announced^[25] and the Sky 3D film The Bachelor King 3D, which Attenborough wrote and narrated, will get a theatrical release in 2012. All Attenborough's 3D projects are being co-produced by Atlantic Productions, as is a second series of First Life, which will explore the origins of the mammals.^[26]

2012 will also herald the third series of David Attenborough's Life Stories for BBC Radio 4 and the new commission David Attenborough's Natural Curiosities on Eden.^[27]

Other work[link]

From 1983, Attenborough worked on two environmentally themed musicals with the WWF and writers Peter Rose and Anne Conlon. Yanomamo was the first, about the Amazon rainforest, and the second, Ocean World, premiered at the Royal Festival Hall in 1991. They were both narrated by Attenborough on their national tour, and recorded on to audio cassette. Ocean World was also filmed for Channel 4 and later released.

In 1990, he highlighted the case of Mahjoub Sharif as part of the BBC's Prisoners of Conscience series.^[28]

In January 2009, the BBC commissioned Attenborough to provide a series of 20 ten-minute monologues covering the history of nature. Entitled David Attenborough's Life Stories, they are broadcast on Radio 4 in the Friday night slot vacated by Alistair Cooke's Letter from America.^[29] Part of Radio 4's A Point of View strand, the talks are also available as podcasts.^[30]

He appeared in the 2009 Children's Prom at the BBC Promenade Concerts and in the Last Night of the Proms on 12 September 2009, playing a floor polisher in Sir Malcolm Arnold's "A Grand, Grand Overture" (after which he was "shot" by Rory Bremner, who was playing the gun).

Attenborough also serves on the advisory board of BBC Wildlife magazine.

Achievements, awards and recognition[link]

Attenborough's contribution to broadcasting and wildlife film-making has brought him international recognition. He has been called "the great communicator, the peerless educator"^[31] and "the greatest broadcaster of our time".^[32] His programmes are often cited as an example of what public service broadcasting should be, even by critics of the BBC, and have influenced a generation of wildlife film-makers.^[33]

Honorary titles[link]

In 2010 it was reported that Attenborough has been given 29 honorary degrees by British universities, more than any other person.^[34] In 1980, he was honoured by the Open University, with whom he has had a close association throughout his career. He also has honorary Doctor of Science awards from the University of Cambridge (1984), the University of Oxford (1988)^[35], the University of Bath (1981)^[36] and the University of St Andrews (2011).^[37] In 2006, the two eldest Attenborough brothers returned to their home city to receive the title of Distinguished Honorary Fellows of the University of Leicester, "in recognition of a record of continuing distinguished service to the University."^[38] David Attenborough was previously awarded an honorary Doctor of Letters degree by the university in 1970, and was made an honorary Freeman of the City of Leicester in 1990.

Attenborough has received the title Honorary Fellow from Clare College, Cambridge (1980), the Zoological Society of London (1998), the Linnean Society (1999), the Institute of Biology (2000) and the Society of Antiquaries (2007).

Recognition[link]

Attenborough has featured as the subject of a number of BBC television programmes. Life on Air (2002) examined the legacy of his work and Attenborough the Controller (2002) focused on his time in charge of BBC Two. He also featured prominently in The Way We Went Wild (2004), a series about natural history television presenters, and 100 Years of Wildlife Films (2007), a special programme marking the centenary of the nature documentary. In 2006, British television viewers were asked to vote for their Favourite Attenborough Moments for a UKTV poll to coincide with the broadcaster's 80th birthday. The winning clip showed Attenborough observing the mimicry skills of the superb lyrebird.

Attenborough was named as the most trusted celebrity in Britain in a 2006 Reader's Digest poll,.^[39] and the following year he won The Culture Show's Living Icon Award.^[40] He has also been named among the 100 Greatest Britons in a 2002 BBC poll and is one of the top ten "Heroes of Our Time" according to New Statesman magazine.^[41]

He has the distinction of having a number of newly discovered species and fossils being named in his honour. In 1993, after discovering that the Mesozoic reptile Plesiosaurus conybeari had not, in fact, been a true plesiosaur, the paleontologist Robert Bakker renamed the species Attenborosaurus conybeari.^[42] A fossilised armoured fish discovered at the Gogo Formation, Western Australia in 2008 was given the name Materpiscis attenboroughi, after Attenborough had filmed at the site and highlighted its scientific importance in Life on Earth.^[43] The Materpiscis fossil is believed to be the earliest organism capable of internal fertilisation.

He has also lent his name to a species of Ecuadorian flowering tree, Blakea attenboroughi, one of the world's largest-pitchered carnivorous plants, Nepenthes attenboroughii,^[44] and one of only four species of long-beaked echidna, the critically endangered Zaglossus attenboroughi, discovered by explorer and zoologist Tim Flannery in the Cyclops Mountains of New Guinea in 1998.

In September 2009, London's Natural History Museum opened the Attenborough Studio, part of its Darwin Centre development.^[45]

Awards[link]

• 1970: BAFTA Desmond Davis Award
• 1974: Commander of the Order of the British Empire (CBE)
• 1980: BAFTA Fellowship
• 1983: Fellow of the Royal Society (FRS)
• 1985: Knighthood^[46]
• 1991: Commander of the Royal Victorian Order (CVO) for producing Queen Elizabeth II's Christmas broadcast for a number of years from 1986
• 1991: Foreign Honorary Member of the American Academy of Arts and Sciences^[47]
• 1996: Companion of Honour (CH) "for services to nature broadcasting"
• 1998: International Cosmos Prize
• 2003: Michael Faraday Prize awarded by the Royal Society
• 2004: Descartes Prize for Outstanding Science Communication Actions
• 2004: Caird Medal of the National Maritime Museum
• 2005: Order of Merit (OM)^[48]
• 2005: Nierenberg Prize for Science in the Public Interest
• 2006: National Outdoor Book Award (Nature and the Environment category), Life in the Underground
• 2006: National Television Awards Special Recognition Award
• 2006: Institute of Ecology and Environmental Management^[49] - Institute Medal in recognition of his outstanding contribution to the public perception and understanding of ecology
• 2006: The Culture Show British Icon Award
• 2007: British Naturalists' Association Peter Scott Memorial Award
• 2009: BAFTA Specialist factual award for Life In Cold Blood
• 2009: Prince of Asturias Award^[50]
• 2010: Fonseca Prize
• 2010: Queensland Museum Medal^[51][52]
•  ???? RSPB Medal^[53][54]
• 2011: Society for the History of Natural History Founders' Medal

Views and advocacy[link]

Environmental causes[link]

Attenborough's programmes have often included references to the impact of human society on the natural world. The last episode of The Living Planet, for example, focuses almost entirely on humans' destruction of the environment and ways that it could be stopped or reversed. Despite this, he has been criticised for not giving enough prominence to environmental messages. Some environmentalists feel that programmes like Attenborough's give a false picture of idyllic wilderness and do not do enough to acknowledge that such areas are increasingly encroached upon by humans.^[55]

However, his closing message from State of the Planet was forthright:

The future of life on earth depends on our ability to take action. Many individuals are doing what they can, but real success can only come if there's a change in our societies and our economics and in our politics. I've been lucky in my lifetime to see some of the greatest spectacles that the natural world has to offer. Surely we have a responsibility to leave for future generations a planet that is healthy, inhabitable by all species.

Attenborough has subsequently become more vocal in his support of environmental causes. In 2005 and 2006, he backed a BirdLife International project to stop the killing of albatross by longline fishing boats.^[56] He gave public support to WWF's campaign to have 220,000 square kilometres of Borneo's rainforest designated a protected area.^[57] He also serves as a vice-president of BTCV, vice-president of Fauna and Flora International, president of Butterfly Conservation and president of Leicestershire and Rutland Wildlife Trust. In 2003 he launched an appeal to create a rainforest reserve in Ecuador in memory of Christopher Parsons, the producer of Life on Earth and a personal friend, who had died the previous year. The same year, he helped to launch ARKive,^[58] a global project instigated by Parsons to gather together natural history media into a digital library. ARKive is an initiative of Wildscreen, of which Attenborough is a patron.^[59] He later became patron of the World Land Trust, and an active supporter. He supported Glyndebourne in their successful application to obtain planning permission for a wind turbine in an Area of Outstanding Natural Beauty, and gave evidence at the planning inquiry arguing in favour of the proposal.

Attenborough has repeatedly said that he considers human overpopulation to be the root cause of many environmental problems. In The Life of Mammals, he made a plea for humans to curb population growth so that other species will not be crowded out. In 2009, he became a patron of Population Matters, (formerly known as the Optimum Population Trust), a UK charity advocating sustainable human populations.

He has written and spoken publicly about the fact that, despite past scepticism, he believes the Earth's climate is warming in a way that is cause for concern, and that this can likely be attributed to human activity.^[60] He summed up his thoughts at the end of his 2006 documentary "Can We Save Planet Earth?" as follows:

In the past, we didn't understand the effect of our actions. Unknowingly, we sowed the wind and now, literally, we are reaping the whirlwind. But we no longer have that excuse: now we do recognise the consequences of our behaviour. Now surely, we must act to reform it — individually and collectively, nationally and internationally — or we doom future generations to catastrophe.

In a 2005 interview with BBC Wildlife magazine, Attenborough said he considered George W. Bush to be the era's top "environmental villain". In 2007, he further elaborated on the USA's consumption of energy in relation to its population. When asked if he thought America to be "the villain of the piece", he responded:

I don't think whole populations are villainous, but Americans are just extraordinarily unaware of all kinds of things. If you live in the middle of that vast continent, with apparently everything your heart could wish for just because you were born there, then why worry? [...] If people lose knowledge, sympathy and understanding of the natural world, they're going to mistreat it and will not ask their politicians to care for it.^[61]

Religion and creationism[link]

In a December 2005 interview with Simon Mayo on BBC Radio Five Live, Attenborough stated that he considers himself an agnostic.^[62] When asked whether his observation of the natural world has given him faith in a creator, he generally responds with some version of this story, making reference to the Loa loa parasitic worm:

My response is that when Creationists talk about God creating every individual species as a separate act, they always instance hummingbirds, or orchids, sunflowers and beautiful things. But I tend to think instead of a parasitic worm that is boring through the eye of a boy sitting on the bank of a river in West Africa, [a worm] that's going to make him blind. And [I ask them], 'Are you telling me that the God you believe in, who you also say is an all-merciful God, who cares for each one of us individually, are you saying that God created this worm that can live in no other way than in an innocent child's eyeball? Because that doesn't seem to me to coincide with a God who's full of mercy'.^[63]

He has explained that he feels the evidence all over the planet clearly shows evolution to be the best way to explain the diversity of life, and that "as far as [he's] concerned, if there is a supreme being then he chose organic evolution as a way of bringing into existence the natural world."

In a BBC Four interview with Mark Lawson, Attenborough was asked if he at any time had any religious faith. He replied simply, "No."^[64] He has also said "It never really occurred to me to believe in God".^[65]

In 2002, Attenborough joined an effort by leading clerics and scientists to oppose the inclusion of creationism in the curriculum of UK state-funded independent schools which receive private sponsorship, such as the Emmanuel Schools Foundation.^[66] In 2009, Attenborough stated that the Book of Genesis, by saying that the world was there for people to dominate, had taught generations that they can "dominate" the environment, and that this has resulted in the devastation of vast areas of the environment.^[67] Attenborough further explained to the science journal Nature, "That's why Darwinism, and the fact of evolution, is of great importance, because it is that attitude which has led to the devastation of so much, and we are in the situation that we are in."^[68][69]

Also in early 2009, the BBC broadcast an Attenborough one-hour special, Charles Darwin and the Tree of Life. In reference to the programme, Attenborough stated that "People write to me that evolution is only a theory. Well, it is not a theory. Evolution is as solid a historical fact as you could conceive. Evidence from every quarter. What is a theory is whether natural selection is the mechanism and the only mechanism. That is a theory. But the historical reality that dinosaurs led to birds and mammals produced whales, that's not theory."^[70] He strongly opposes creationism and its offshoot "intelligent design", saying that a survey that found a quarter of science teachers in state schools believe that creationism should be taught alongside evolution in science lessons was "really terrible".^[70]

In March 2009, Attenborough appeared on Friday Night with Jonathan Ross. Attenborough stated that he felt evolution did not rule out the existence of a God and accepted the title of agnostic saying, "My view is: I don't know one way or the other but I don't think that evolution is against a belief in God."^[71]

Attenborough has joined evolutionary biologist Richard Dawkins and other top scientists in signing a campaign statement coordinated by the British Humanist Association (BHA). The statement calls for "creationism to be banned from the school science curriculum and for evolution to be taught more widely in schools."^[72]

BBC and public service broadcasting[link]

Attenborough is a lifelong supporter of the BBC, public broadcasting and the television licence. He has said:

PSB, to me, is not about selecting individual programme strands here or there, financing them from some outside source and then foisting them upon commercial networks. Public Service Broadcasting, watched by a healthy number of viewers, with programmes financed in proportion to their intrinsic needs and not the size of the audience, can only effectively operate as a network — a network whose aim is to cater for the broadest possible range of interests, popular as well as less popular, a network that measures its success not only by its audience size but by the range of its schedule.^[73]

Public service broadcasting is one of the things that distinguishes this country and makes me want to live here. I have spent all my life in it. I would be very distressed if public service broadcasting was weakened. I have been at the BBC since 1952, and know the BBC is constantly being battered. It is today.^[74]

If you could demonstrate that the BBC was grossly extravagant there might be a case for saying OK take it away. But in fact the BBC per minute in almost every category is as cheap as you can find anywhere in the world and produces the best quality. If you take the money away, which part of the BBC will you remove? The BBC has gone through swingeing staff cuts. It has been cut to the bone, if you divert licence fee money elsewhere, you cut quality and services. There is always that threat from politicians who will say your licence fee is up for grabs. We will take it. There is a lot of people who want to see the BBC weakened. They talk of this terrible tax of the licence fee. Yet it is the best bargain that is going. Four radio channels and god knows how many TV channels. It is piffling.^[74]

There have always been politicians or business people who have wanted to cut the BBC back or stop it saying the sort of things it says. There's always been trouble about the licence and if you dropped your guard you could bet our bottom dollar there'd be plenty of people who'd want to take it away. The licence fee is the basis on which the BBC is based and if you destroy it, broadcasting... becomes a wasteland.^[75]

Attenborough expressed regret at some of the changes made to the BBC in the 1990s by Director-General John Birt, who introduced an internal market at the corporation, slimmed and even closed some departments and outsourced much of the corporation’s output to private production companies, in line with the Broadcasting Act 1990. He has said:

There is no question but that Birtism . . . has had some terrible results. On the other hand, the BBC had to change. Now it has to produce programmes no one else can do. Otherwise, forget the licence fee.^[76]

The Bristol Unit has suffered along with the rest of the BBC from recent staff cuts. Yet it remains confident in the belief that the BBC will maintain it, in spite of the vagaries of fashion, because the Corporation believes that such programmes deserve a place in the schedules of any broadcaster with pretensions of providing a Public Service. In due course, similar specialist Units were also established in London, in order to produce programmes on archaeology and history, on the arts, on music and on science. They too, at one time, had their successes. But they have not survived as well as the Unit in Bristol. The statutory requirement that a certain percentage of programmes must come from independent producers has reduced in-house production and the Units necessarily shrank proportionately in size. As they dwindled, so the critical mass of their production expertise has diminished. The continuity of their archives has been broken, they have lost the close touch they once had worldwide with their subjects and they are no longer regarded internationally as the centres of innovation and expertise that they once were.^[77]

When Birt gets up and says the whole of the BBC was a creative mess and it was wasteful, I never saw any evidence of that. I absolutely know it wasn’t so in my time. Producers now spend all their time worrying about money, and the thing has suffered for it.^[78]

In 2008, he criticised the BBC’s television schedules:

I have to say that there are moments when I wonder — moments when its two senior networks, first set up as a partnership, schedule simultaneously programmes of identical character, thereby contradicting the very reason that the BBC was given a second network. Then there are times when both BBC One and BBC Two, intoxicated by the sudden popularity of a programme genre, allow that genre to proliferate and run rampant through the schedules. The result is that other kinds of programmes are not placed, simply because of a lack of space. Do we really require so many gardening programmes, make-over programmes or celebrity chefs? Is it not a scandal in this day and age, that there seems to be no place for continuing series of programmes about science or serious music or thoughtful in-depth interviews with people other than politicians?^[73]

In 2009, Attenborough commented on the general state of British television, describing the newly introduced product placement on commercial television as something he considered an "appalling" idea 20 years earlier:

I think it's in great trouble. The whole system on which it was built — a limited number of networks, with adequate funding — is under threat. That funding is no longer there. As stations proliferate, so audiences are reduced. The struggle for audiences becomes ever greater, while money diminishes. I think that's a fair recipe for trouble. Inevitably, this has an impact on the BBC … Fortunately, the BBC doesn't think natural history programmes must compete with Strictly Come Dancing in terms of audience. The BBC says, 'Make proper, responsible natural history programmes.'^[79]

Other causes[link]

In May 2005, Attenborough was appointed as patron of the UK's Blood Pressure Association, which provides information and support to people with hypertension.^[80]

Attenborough is also an honorary member of BSES Expeditions, a youth development charity that operates challenging scientific research expeditions to remote wilderness environments.

Parodies and artistic portrayals[link]

Attenborough's accent and hushed, excited delivery have been the subject of frequent parodies by comedians, most notably Spike Milligan, Marty Feldman and The Goodies. Attenborough is portrayed by Michael Palin in the final episode of Monty Python's Flying Circus, where he searches the African jungle for the legendary Walking Tree of Dahomey (Quercus Nicholas Parsonus), sweating excessively and accompanied by native guides wearing saxophones. David Attenborough has also been parodied in a series of GEICO insurance commercials, showing a nature show host, clearly patterned on Attenborough.

"Time Flies", a sketch by David Ives, features a pair of anthropomorphic mayflies engaging in a courtship ritual, while watching themselves on television in a documentary narrated by David Attenborough.^[81]

The character of Nigel Thornberry, a nature documentarian on Nickelodeon's The Wild Thornberrys is strongly influenced by Attenborough.

He has also been parodied by the Australia 1980s sketch show The Comedy Company where Ian McFadyen portrays a character called David Rabbitborough, who goes about examining the "wildlife" of various Australian urban stereotypes.

The video game Discworld, based on the series of books by Terry Pratchett, parodies his unique delivery to explain different aspects of the Discworld Universe, such as L-Space.

Mythbusters host Adam Savage often imitates Attenborough when speaking about his co-host Jamie Hyneman, which Jamie verified in the YouTube Special Episode.

Filmography[link]

Attenborough is known foremost for writing and presenting the ten Life series, which are presented in chronological order below:

• Life on Earth (1979)
• The Living Planet (1984)
• The Trials of Life (1990)
• Life in the Freezer (1993)
• The Private Life of Plants (1995)
• The Life of Birds (1998)
• The Life of Mammals (2002)
• Life in the Undergrowth (2005)
• Life in Cold Blood (2008)
• First Life (2010)

His voice is synonymous with wildlife documentaries for British audiences, and the principal series with which his narration is associated include:

• Wildlife on One (1977–2005)
• BBC Wildlife Specials (1995–2008)
• The Blue Planet (2001)
• Planet Earth (British version) (2006)
• Nature's Great Events (2009)
• Life (2009)
• Frozen Planet (2011)

Books[link]

Bibliography[link]

• Zoo Quest to Guyana (Lutterworth Press, 1956)
• Zoo Quest for a Dragon (Lutterworth Press, 1957)
  • (book club edition with 85 extra pages, Quest for the Paradise Birds, 1959)
• Zoo Quest in Paraguay (Lutterworth Press, 1959)
  • paperback (Penguin Books, 1982)
• Quest in Paradise (1960)
• People of Paradise (Harper & Brothers, 1960)
• Zoo Quest to Madagascar (1961)
• Quest Under Capricorn (1963)
• Fabulous Animals (BBC, 1975) ISBN 0-563-17006-9
• The Tribal Eye (1976)
• Life on Earth (1979)
• Discovering Life on Earth (1981)
• Journeys to the past: Travels in New Guinea, Madagascar, and the northern territory of Australia (1983) Penguin Books ISBN 0-14-00.64133
• The Living Planet (1984)
• The First Eden: The Mediterranean World and Man (1987)(Little Brown & Co (T); 1st American ed edition (March 1990))
• The Atlas of the Living World (1989)
• The Trials of Life (Collins, 1990) ISBN 0-00-219912-2
• The Private Life of Plants (BBC Books, 1994) ISBN 0-563-37023-8
• The Life of Birds (BBC Books, 1998) ISBN 0-563-38792-0
• The Life of Mammals (BBC Books, 2002) ISBN 0-563-53423-0
• Life on Air: Memoirs of a Broadcaster (autobiography; 2002) ISBN 0-563-53461-3
  • paperback: ISBN 0-563-48780-1
  • updated edition: ISBN 978-1-84607-652-7
• Life in the Undergrowth (BBC Books, 2005) ISBN 0-563-52208-9
• Amazing Rare Things - The Art of Natural History in the Age of Discovery with Susan Owens, Martin Clayton and Rea Alexandratos (The Royal Collection, 2007) Hardback - ISBN 978-1-902163-46-8; Softback - ISBN 978-1-902163-99-4
• Life in Cold Blood (BBC Books, 2007) ISBN 978-0-563-53922-3
• Life Stories, (HarperCollins Publishers, 2009) ISBN 978-0-00-733883-2
• New Life Stories, (Collins, 2011) ISBN 978-0-00-742512-9

Introductions[link]

Attenborough has written the introduction or foreword for a number of books, including:

• Buxton, Cindy; Annie Price (1983). Survival South Atlantic. London :: Granada,. pp. xiii,237p.,[64]p. of plates : ill(some col.), maps,col.ports. ; 26cm.. ISBN 0-246-12087-8. 
• African Jigsaw: A Musical Entertainment, Peter Rose and Anne Conlon (published: 1986, Weinberger)
• Tomorrow Is Too Late, Various (The Macmillan Press, 1990)
• Life in the Freezer: Natural History of the Antarctic, Alastair Fothergill (BBC Books, 1993), ISBN 0-563-36431-9
• Birds of Paradise: Paradisaeidae (Bird Families of the World series) Clifford B. Frith, Bruce M. Beehler, William T. Cooper (Illustrator) (Oxford University Press, 1998) ISBN 0-19-854853-2
• The Blue Planet, Andrew Byatt, Alastair Fothergill, Martha Holmes (BBC Books, 2001) ISBN 0-563-38498-0.
• Light on the Earth (BBC Books, 2005), two decades of winning images from the Wildlife Photographer of the Year competition, ISBN 0-563-52260-7
• Planet Earth, Alastair Fothergill (BBC Books, 2006), ISBN 0-563-52212-7

Audio recordings[link]

• Tarka the Otter by Henry Williamson (available on audiocassette, 1978)
• Yanomamo (musical entertainment, 1983) by Peter Rose and Anne Conlon; on-stage narration and published audio recording
• Ocean World (musical entertainment, 1990) by Peter Rose and Anne Conlon; on-stage narration (including at The Royal Festival Hall), for audio recording and video broadcast (both published)
• Peter and the Wolf for BBC Music Magazine (free CD with the June 2000 issue).

In addition, Attenborough has recorded some of his own works in audiobook form, including Life on Earth, Zoo Quest for a Dragon and his autobiography Life on Air: Memoirs of a Broadcaster.

References[link]

External links[link]

Wikiquote has a collection of quotations related to: David Attenborough

Wikimedia Commons has media related to: David Attenborough

• David Attenborough at the Internet Movie Database
• BBC Books David Attenborough website
• BBC Life on Air website
• BBC biography
• Wildfilmhistory.org biography
• BBC interviews with Attenborough in 1976 and 1998
• PBS interview with Attenborough in 1998
• A Life in TV: Sir David Attenborough, BAFTA
• BBC Wildlife Finder - David Attenborough's favourite moments
• Tribute from the World Land Trust
• David Attenborough profile on Eden Channel website
• List of David Attenborough's programmes on Eden Channel website