biography information definition

Definition of Biography

A biography is the non- fiction , written history or account of a person’s life. Biographies are intended to give an objective portrayal of a person, written in the third person. Biographers collect information from the subject (if he/she is available), acquaintances of the subject, or in researching other sources such as reference material, experts, records, diaries, interviews, etc. Most biographers intend to present the life story of a person and establish the context of their story for the reader, whether in terms of history and/or the present day. In turn, the reader can be reasonably assured that the information presented about the biographical subject is as true and authentic as possible.

Biographies can be written about a person at any time, no matter if they are living or dead. However, there are limitations to biography as a literary device. Even if the subject is involved in the biographical process, the biographer is restricted in terms of access to the subject’s thoughts or feelings.

Biographical works typically include details of significant events that shape the life of the subject as well as information about their childhood, education, career, and relationships. Occasionally, a biography is made into another form of art such as a film or dramatic production. The musical production of “Hamilton” is an excellent example of a biographical work that has been turned into one of the most popular musical productions in Broadway history.

Common Examples of Biographical Subjects

Most people assume that the subject of a biography must be a person who is famous in some way. However, that’s not always the case. In general, biographical subjects tend to be interesting people who have pioneered something in their field of expertise or done something extraordinary for humanity. In addition, biographical subjects can be people who have experienced something unusual or heartbreaking, committed terrible acts, or who are especially gifted and/or talented.

As a literary device, biography is important because it allows readers to learn about someone’s story and history. This can be enlightening, inspiring, and meaningful in creating connections. Here are some common examples of biographical subjects:

• political leaders
• entrepreneurs
• historical figures
• serial killers
• notorious people
• political activists
• adventurers/explorers
• religious leaders
• military leaders
• cultural figures

Famous Examples of Biographical Works

The readership for biography tends to be those who enjoy learning about a certain person’s life or overall field related to the person. In addition, some readers enjoy the literary form of biography independent of the subject. Some biographical works become well-known due to either the person’s story or the way the work is written, gaining a readership of people who may not otherwise choose to read biography or are unfamiliar with its form.

Here are some famous examples of biographical works that are familiar to many readers outside of biography fans:

• Alexander Hamilton (Ron Chernow)
• Prairie Fires: The American Dreams of Laura Ingalls Wilder (Caroline Fraser)
• Steve Jobs (Walter Isaacson)
• Churchill: A Life (Martin Gilbert)
• The Professor and the Madman: A Tale of Murder, Insanity, and the Making of the Oxford English Dictionary (Simon Winchester)
• A Beautiful Mind (Sylvia Nasar)
• The Black Rose (Tananarive Due)
• John Adams (David McCullough)
• Into the Wild ( Jon Krakauer )
• John Brown (W.E.B. Du Bois)
• Frida: A Biography of Frida Kahlo (Hayden Herrera)
• The Immortal Life of Henrietta Lacks (Rebecca Skloot)
• Team of Rivals: The Political Genius of Abraham Lincoln (Doris Kearns Goodwin)
• Shirley Jackson : A Rather Haunted Life ( Ruth Franklin)
• the stranger in the Woods: The Extraordinary Story of the Last True Hermit (Michael Finkel)

Difference Between Biography, Autobiography, and Memoir

Biography, autobiography , and memoir are the three main forms used to tell the story of a person’s life. Though there are similarities between these forms, they have distinct differences in terms of the writing, style , and purpose.

A biography is an informational narrative and account of the life history of an individual person, written by someone who is not the subject of the biography. An autobiography is the story of an individual’s life, written by that individual. In general, an autobiography is presented chronologically with a focus on key events in the person’s life. Since the writer is the subject of an autobiography, it’s written in the first person and considered more subjective than objective, like a biography. In addition, autobiographies are often written late in the person’s life to present their life experiences, challenges, achievements, viewpoints, etc., across time.

Memoir refers to a written collection of a person’s significant memories, written by that person. Memoir doesn’t generally include biographical information or chronological events unless it’s relevant to the story being presented. The purpose of memoir is reflection and an intention to share a meaningful story as a means of creating an emotional connection with the reader. Memoirs are often presented in a narrative style that is both entertaining and thought-provoking.

Examples of Biography in Literature

An important subset of biography is literary biography. A literary biography applies biographical study and form to the lives of artists and writers. This poses some complications for writers of literary biographies in that they must balance the representation of the biographical subject, the artist or writer, as well as aspects of the subject’s literary works. This balance can be difficult to achieve in terms of judicious interpretation of biographical elements within an author’s literary work and consideration of the separate spheres of the artist and their art.

Literary biographies of artists and writers are among some of the most interesting biographical works. These biographies can also be very influential for readers, not only in terms of understanding the artist or writer’s personal story but the context of their work or literature as well. Here are some examples of well-known literary biographies:

Example 1:  Savage Beauty: The Life of Edna St. Vincent Millay  (Nancy Milford)

One of the first things Vincent explained to Norma was that there was a certain freedom of language in the Village that mustn’t shock her. It wasn’t vulgar. ‘So we sat darning socks on Waverly Place and practiced the use of profanity as we stitched. Needle in, . Needle out, piss. Needle in, . Needle out, c. Until we were easy with the words.’

This passage reflects the way in which Milford is able to characterize St. Vincent Millay as a person interacting with her sister. Even avid readers of a writer’s work are often unaware of the artist’s private and personal natures, separate from their literature and art. Milford reflects the balance required on the part of a literary biographer of telling the writer’s life story without undermining or interfering with the meaning and understanding of the literature produced by the writer. Though biographical information can provide some influence and context for a writer’s literary subjects, style, and choices , there is a distinction between the fictional world created by a writer and the writer’s “real” world. However, a literary biographer can illuminate the writer’s story so that the reader of both the biography and the biographical subject’s literature finds greater meaning and significance.

Example 2:  The Invisible Woman: The Story of Nelly Ternan and Charles Dickens  (Claire Tomalin)

The season of domestic goodwill and festivity must have posed a problem to all good Victorian family men with more than one family to take care of, particularly when there were two lots of children to receive the demonstrations of paternal love.

Tomalin’s literary biography of Charles Dickens reveals the writer’s extramarital relationship with a woman named Nelly Ternan. Tomalin presents the complications that resulted for Dickens from this relationship in terms of his personal and family life as well as his professional writing and literary work. Revealing information such as an extramarital relationship can influence the way a reader may feel about the subject as a person, and in the case of literary biography it can influence the way readers feel about the subject’s literature as well. Artists and writers who are beloved , such as Charles Dickens, are often idealized by their devoted readers and society itself. However, as Tomalin’s biography of Dickens indicates, artists and writers are complicated and as subject to human failings as anyone else.

Example 3:  Virginia Woolf  (Hermione Lee)

‘A self that goes on changing is a self that goes on living’: so too with the biography of that self. And just as lives don’t stay still, so life-writing can’t be fixed and finalised. Our ideas are shifting about what can be said, our knowledge of human character is changing. The biographer has to pioneer, going ‘ahead of the rest of us, like the miner’s canary, testing the atmosphere , detecting falsity, unreality, and the presence of obsolete conventions’. So, ‘There are some stories which have to be retold by each generation’. She is talking about the story of Shelley, but she could be talking about her own life-story.

In this passage, Lee is able to demonstrate what her biographical subject, Virginia Woolf, felt about biography and a person telling their own or another person’s story. Literary biographies of well-known writers can be especially difficult to navigate in that both the author and biographical subject are writers, but completely separate and different people. As referenced in this passage by Lee, Woolf was aware of the subtleties and fluidity present in a person’s life which can be difficult to judiciously and effectively relay to a reader on the part of a biographer. In addition, Woolf offers insight into the fact that biographers must make choices in terms of what information is presented to the reader and the context in which it is offered, making them a “miner’s canary” as to how history will view and remember the biographical subject.

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Biographical Information

What is a biography, general suggestion for finding biographical information, general sources - biography databases, general sources - reference books, biographical master indexes.

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• A biography is an account of the special and important events in a person's life
• Not to be confused with bibliography , which is a listing of books and articles on a topic
• Biographies may be brief and cover only basic information about a person's life such as dates of birth and death, education and vocation
• A biography may also be very detailed, and cover the cultural background, outstanding accomplishments, and historical significance of an individual.
• Biographical sources cover living and deceased persons, notable persons in particular countries, persons in specific occupations, celebrities, and civil and government leaders
• The person's full name and correct spelling (for example: Smith, Smyth or Smythe)
• date of birth
• date of death (when applicable)
• nationality or country of residence
• occupation or profession

For basic facts about a person (e.g.,"When was Napoleon born?"):

• General encyclopedias

Can’t find biographical information in general encyclopedias or need more information? Check:

• One of the general sources listed on this page
• The specialized biographical reference books listed on the other tabs in this guide

Can’t find the person you are looking for or get enough information from a biographical reference book? Check:

• One of the biographical master indexes listed on this page. These guide you to books, periodical articles or other reference sources. Please note that sometimes you will be referred to another index.

For highly detailed information :

• Search UToledo’s online library catalog for books on the person:
• Perform a subject search by typing the person's last name followed by their first name, e.g., Whitman, Walt
• Execute the search and look for the subheading --Biography. The complete heading in this example is: Whitman, Walt, 1819-1892 – Biography
• You can also use the UToledo catalog to search for the titles of books and periodicals found in the biographical master indexes

If UToledo does not own a title you want :

• Use the OhioLINK Library Catalog to directly borrow books from other college libraries in Ohio. Books arrive within three to five business days
• There is generally no charge for this service
• The library may need two to three weeks to get the item you need
• For assistance with the library catalog, OhioLINK, or interlibrary loan forms, consult a reference librarian at the Information/Reference Desk.

Biographical information may not be available for all individuals. In these cases:

• Information about an author can sometimes be found by checking the preface or introduction of an author's work for scholarly background and academic achievements
• Perform an author search in the library catalog by typing in the last name followed by the first name

Master Indexes provide a name index to books, periodical articles, and other references where you might find a biography or obituary of someone.  Usually, you can search by name (last, first) and then be given a code for a title, volume, and page number.  Consult the key at the beginning of the book to decipher the title you need.  Search by Title in the UToledo Library Catalog to see if we have the book or periodical, or order it from another OhioLINK library.

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Example sentences biographical information

Definition of 'biographical' biographical.

Definition of 'information' information

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[ bahy- og -r uh -fee , bee- ]

the biography of Byron by Marchand.

• an account in biographical form of an organization, society, theater, animal, etc.
• such writings collectively.
• the writing of biography as an occupation or field of endeavor.

/ baɪˈɒɡrəfɪ; ˌbaɪəˈɡræfɪkəl /

• an account of a person's life by another
• such accounts collectively
• The story of someone's life. The Life of Samuel Johnson , by James Boswell , and Abraham Lincoln , by Carl Sandburg , are two noted biographies. The story of the writer's own life is an autobiography .

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Derived forms.

• biˈographer , noun
• biographical , adjective
• ˌbioˈgraphically , adverb

Word History and Origins

Origin of biography 1

Example Sentences

Barrett didn’t say anything on Tuesday to contradict our understanding of her ideological leanings based on her past rulings, past statements and biography.

Republicans, meanwhile, focused mostly on her biography — including her role as a working mother of seven and her Catholic faith — and her credentials, while offering few specifics about her record as a law professor and judge.

She delivered an inspiring biography at one point, reflecting on the sacrifice her mother made to emigrate to the United States.

As Walter Isaacson pointed out in his biography of Benjamin Franklin, Franklin proposed the postal system as a vital network to bond together the 13 disparate colonies.

Serving that end, the book is not an in-depth biography as much as a summary of Galileo’s life and science, plus a thorough recounting of the events leading up to his famous trial.

The Amazon biography for an author named Papa Faal mentions both Gambia and lists a military record that matches the FBI report.

For those unfamiliar with Michals, an annotated biography and useful essays are included.

Did you envision your Pryor biography as extending your previous investigation—aesthetically and historically?

But Stephen Kotkin's new biography reveals a learned despot who acted cunningly to take advantage of the times.

Watching novelists insult one another is one of the primary pleasures of his biography.

He also published two volumes of American Biography, a work which his death abridged.

Mme. de Chaulieu gave her husband the three children designated in the duc's biography.

The biography of great men always has been, and always will be read with interest and profit.

I like biography far better than fiction myself: fiction is too free.

The Bookman: "A more entertaining narrative whether in biography or fiction has not appeared in recent years."

Related Words

• autobiography

Libraries | Research Guides

Biographical information.

• Biographical Sources
• Searching for Biographical Sources in NUsearch

Biographical resources

• Online resources
• Print resources
• American National Biography This link opens in a new window There is no need to sign in at website to access this resource. Begin searching or browsing for articles. The American National Biography is the premier, authoritative, historical biographical encyclopedia for the United States. Over 18,700 men and women are included, and the resource is updated quarterly with new entries and revisions of previously published entries to enhance their accuracy and currency. Articles are by established scholars. Living individuals are not included.
• Oxford Dictionary of National Biography This link opens in a new window The Oxford Dictionary of National Biography provides biographies of over 50,000 individuals worldwide who have shaped the history of the British Isles. Access restricted to 3 simultaneous users at a time.
• Gale in Context: Biography This link opens in a new window Gale In Context: Biography is an engaging online experience for those seeking contextual information on the world's most influential people. Biography merges Gale's authoritative reference content with periodicals and multimedia organized into a user-friendly portal experience while allowing users to search for people based on name, occupation, nationality, ethnicity, birth/death dates and places, or gender as well as keyword and full text.
• Current Biography Illustrated This link opens in a new window Current Biography Illustrated contains profiles of individuals living at the time of publication, often at the height of, or even the beginning of, their fame or notoriety. Occasionally a revised biography will be published, and a brief obituary, but published biographies are never revised in light of later events, so a useful source for seeing how individuals were perceived at the time they were active. Illustrations for most individuals are included.
• Chambers Biographical Dictionary International and historical coverage of all areas of human achievement including the arts, science, technology, sport, politics, philosophy and business.
• World Biographical Information System WBIS This link opens in a new window Over 3.6 million short biographical entries for individuals who lived across the world from ancient times to the present. Includes full text images of the original sources, usually older biographical dictionaries. Especially valuable for locating information on hard-to-find individuals from the past.
• Almanac of American Politics Includes profiles of every member of Congress and every governor. It offers in-depth and completely up-to-date narrative profiles of all 50 states and 435 House districts.
• African American Biographical Database This link opens in a new window From July 1, 2023 AABD became a component of ProQuest Black Studies Center. Biographical sketches and essays on African Americans from 1790 to 1950. The Database corresponds to the printed Black Biographical Dictionaries. These sketches have been assembled from biographical dictionaries, yearbooks, directories, histories, personal accounts, and other published sources. The full text of 300 rare books is displayed and searchable in page images, intact with images and illustrations.
• Complete Dictionary of Scientific Biography Provides information on the history of science through articles on the professional lives of scientists. Covers all periods of science from classical antiquity to modern times.
• Gale Literature Resource Center This link opens in a new window Provides access to biographies, bibliographies, and critical analayses of authors from every age and literary discipline. Covers more than 120,000 novelists, essayist, poets, journalists, and other writers, with in-depth coverage of 2,500 of the most-studied authors. Includes links to: Dictionary of literary biography, Contemporary authors, Contemporary literary criticism, and more.
• Palgrave MacMillan Dictionary of Women’s Biography Biographical information about women: historical, current, and worldwide.

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• Last Updated: Jul 7, 2023 1:49 PM
• URL: https://libguides.northwestern.edu/biographical

Biographical Resources: A Research Guide: Introduction

Introduction.

• National and International Biographies
• Biographical Indexes
• K. G. Saur Indexes & Microfiche
• Subject Biographies
• Dissertations and Theses
• Research Help

We purchase access to new online versions of major biographical reference sources as they become available. Many important biographical resources are available in print and on microfilm. This guide combines online titles with the selected microform and print biography titles in the Olin and Africana reference collections. Online databases are available to Cornell users only.

Biography is a branch of the study of history. The reliability of biographical sources varies widely and is subject to the usual vagaries of historical studies: lack of accurate information, too much or conflicting information, too little information, psychological theorizing, etc. But a well-written biographical article in a reliable reference book or database can be a source of both pleasure and enlightenment. Enjoy!

For further information or to locate titles not listed here, always feel free to consult with the reference staff .

Reference Help

Permissions Information

If you wish to use or adapt any or all of the content of this Guide go to Cornell Library's Research Guides Use Conditions to review our use permissions and our Creative Commons license.

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• Last Updated: Apr 25, 2024 4:28 PM
• URL: https://guides.library.cornell.edu/biographyresearch

MERRIMACK COLLEGE MCQUADE LIBRARY

General reference.

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What is Biographical Information?

Library databases.

• Books & Publishing
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• Cities, States, Regions
• Colleges & Universities
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• Economy / Finance
• Elements / Periodic Table
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• Images & Audiovisual
• Information Literacy
• Internet & the WWW
• Maps & Gazetteers
• Movie Reviews
• Public Opinion Polls
• Primary Sources
• Study Guides

Biographical information is information about a particular person's life including

• date of birth
• date of death
• accomplishments
• occupations

Find biographies of people, searching by occupation, nationality, ethnicity, birth and death date, and of course by name.  

Search hundreds of reference sources - dictionaries, bilingual dictionaries, thesauri, encyclopedias, quotations and atlases - for topic overviews and links to our other online resources. Includes mind map/concept map search feature.

• Gale Ebooks - Biography Explore Biography Reference sources in the Gale Ebooks Collection.
• Nexis Uni This link opens in a new window Use the Research People box to search for biographical information for national and international persons.

Note:  Access to library databases is restricted to current student , faculty and staff ONLY .

• Biography.com
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What Is Biography? Definition, Usage, and Literary Examples

Biography definition.

A  biography  (BYE-og-ruh-fee) is a written account of one person’s life authored by another person. A biography includes all pertinent details from the subject’s life, typically arranged in a chronological order. The word  biography  stems from the Latin  biographia , which succinctly explains the word’s definition:  bios  = “life” +  graphia  = “write.”

Since the advent of the written word, historical writings have offered information about real people, but it wasn’t until the 18th century that biographies evolved into a separate literary genre.  Autobiographies  and memoirs fall under the broader biography genre, but they are distinct literary forms due to one key factor: the subjects themselves write these works. Biographies are popular source materials for documentaries, television shows, and motion pictures.

The History of Biographies

The biography form has its roots in Ancient Rome and Greece. In 44 BCE, Roman writer Cornelius Nepos published  Excellentium Imperatorum Vitae  ( Lives of the Generals ), one of the earliest recorded biographies. In 80 CE, Greek writer Plutarch released  Parallel Lives , a sweeping work consisting of 48 biographies of famous men. In 121 CE, Roman historian Suetonius wrote  De vita Caesarum  ( On the Lives of the Caesars ), a series of 12 biographies detailing the lives of Julius Caesar and the first 11 emperors of the Roman Empire. These were among the most widely read biographies of their time, and at least portions of them have survived intact over the millennia.

During the Middle Ages, the Roman Catholic Church had a notable influence on biographies. Historical, political, and cultural biographies fell out of favor. Biographies of religious figures—including saints, popes, and church founders—replaced them. One notable exception was Italian painter/architect Giorgio Vasari’s 1550 biography,  The Lives of the Most Excellent Painters, Sculptors, and Architects , which was immensely popular. In fact, it is one of the first examples of a bestselling book.

Still, it wasn’t until the 18th century that authors began to abandon multiple subjects in a single work and instead focus their research and writing on one subject. Scholars consider James Boswell’s 1791  The Life of Samuel Johnson  to be the first modern biography. From here, biographies were established as a distinct literary genre, separate from more general historical writing.

As understanding of psychology and sociology grew in the 19th and early 20th centuries, biographies further evolved, offering up even more comprehensive pictures of their subjects. Authors who played major roles in this contemporary approach to biographing include Lytton Strachey, Gamaliel Bradford, and Robert Graves.

Types of Biographies

While all biographical works chronicle the lives of real people, writers can present the information in several different ways.

• Popular biographies are life histories written for a general readership.  The Immortal Life of Henrietta Lacks  by Rebecca Skloot and  Into the Wild  by Jon Krakauer are two popular examples.
• Critical biographies discuss the relationship between the subject’s life and the work they produced or were involved in; for example,  The Billionaire Who Wasn’t: How Chuck Feeney Secretly Made and Gave Away a Fortune  by Conor O’Clery and  Unpresidented: A Biography of Donald Trump  by Martha Brockenbrough.
• Historical biographies put greater understanding on how the subject’s life and contributions affected or were affected by the times in which they lived; see  John Adams  by David McCullough and  Catherine the Great  by Peter K. Massie.
• Literary biographies concentrate almost exclusively on writers and artists, blending a conventional  narrative  of the historical facts of the subject’s life with an exploration of how these facts impacted their creative output. Some examples include  Savage Beauty: The Life of Edna St. Vincent Millay  by Nancy Milford and  Jackson Pollock: An American Saga  by Gregory White Smith and Steven Naifeh.
• Reference biographies are more scholarly writings, usually written by multiple authors and covering multiple lives around a single topic. They verify facts, provide background details, and contribute supplemental information resources, like bibliographies, glossaries, and historical documents; for example,  Black Americans in Congress, 1870-2007  and the  Dictionary of Canadian Biography .
• Fictional biographies, or biographical novels, like  The Other Boleyn Girl  by Philippa Gregory, incorporate creative license into the retelling of a real person’s story by taking on the structure and freedoms of a novel. The term can also describe novels in which authors give an abundance of background information on their characters, to the extent that the novel reads more like a biography than fiction. An example of this is George R.R. Martin’s  Fire and Blood , a novel detailing the history of a royal family from his popular  A Song of Ice and Fire

Biographies and Filmed Entertainment

Movie makers and television creators frequently produce biographical stories, either as dramatized productions based on real people or as nonfiction accounts.

Documentary

This genre is a nonfictional movie or television show that uses historical records to tell the story of a subject. The subject might be a one person or a group of people, or it might be a certain topic or theme. To present a biography in a visually compelling way, documentaries utilize archival footage, recreations, and interviews with subjects, scholars, experts, and others associated with the subject.

Famous film documentaries include  Grey Gardens,  a biography of two of Jacqueline Kennedy’s once-wealthy cousins, who, at the time of filming, lived in squalor in a condemned mansion in the Hamptons; and  I Am Not Your Negro , a biography of the life and legacy of pioneering American author James Baldwin.

Television documentary series tell one story over the course of several episodes, like  The Jinx :  The Life and Deaths of Robert Durst , a biography of the real estate heir and alleged serial killer that focused on his suspected crimes. There are many nonfiction television shows that use a documentary format, but subjects typically change from one episode to the next, such as A&E’s  Biography  and PBS’s  POV .

These films are biographical motion pictures, written by screenwriters and performed by actors. They often employ a certain amount of creative liberty in their interpretation of a real life. This is largely done to maintain a feasible runtime; capturing all of the pivotal moments of a subject’s life in a 90- or 120-minute movie is all but impossible. So, filmmakers might choose to add, eliminate, or combine key events and characters, or they may focus primarily on one or only a few aspects of the subject’s life. Some popular examples:  Coal Miner’s Daughter , a biography of country music legend Loretta Lynn;  Malcom X , a biopic centered on the civil rights leader of the same name; and  The King’s Speech , a dramatization of Prince Albert’s efforts to overcome a stutter and ascend the English throne.

Semi-fictionalized account

This approach takes a real-life event and interprets or expands it in ways that stray beyond what actually happened. This is done for entertainment and to build the story so it fits the filmmaker’s vision or evolves into a longer form, such as a multi-season television show. These accounts sometimes come with the disclaimer that they are “inspired by true events.” Examples of semi-fictionalized accounts are the TV series  Orange Is the New Black ,  Masters of Sex , and  Mozart of the Jungle —each of which stem from at least one biographical element, but showrunners expounded upon to provide many seasons of entertainment.

The Functions of Biography

Biographies inform readers about the life of a notable person. They are a way to introduce readers to the work’s subject—the historical details, the subject’s motivations and psychological underpinnings, and their environment and the impact they had, both in the short and long term.

Because the author is somewhat removed from their subject, they can offer a more omniscient, third-person narrative account. This vantage point allows the author to put certain events into a larger context; compare and contrast events, people, and behaviors predominant in the subject’s life; and delve into psychological and sociological themes of which the subject may not have been aware.

Also, a writer structures a biography to make the life of the subject interesting and readable. Most biographers want to entertain as well as inform, so they typically use a traditional  plot  structure—an introduction,  conflict , rising of tension, a climax, a resolution, and an ending—to give the life story a narrative shape. While the ebb and flow of life is a normal day-to-day rhythm, it doesn’t necessarily make for entertaining reading. The job of the writer, then, becomes one of shaping the life to fit the elements of a good plot.

Writers Known for Biographies

Many modern writers have dedicated much of their careers to biographies, such as:

• Kitty Kelley, author of  Jackie Oh! An Intimate Biography; His Way: The Unauthorized Biography of Frank Sinatra ; and  The Family: The Real Story of the Bush Dynasty
• Antonia Fraser, author of  Mary Queen of Scots ;  Cromwell; Our Chief of Men ; and  The Gunpowder Plot: Terror and Faith in 1605
• David McCullough, author of  The Path Between the Seas; Truman ; and  John Adams
• Andrew Morton, author of  Diana: Her True Story in Her Own Words; Madonna ; and  Tom Cruise: An Unauthorized Biography
• Alison Weir, author of  The Six Wives of Henry VIII; Eleanor of Aquitaine: By the Wrath of God; Queen of England ; and  Katherine Swynford: The Story of John of Gaunt and His Scandalous Duchess

Examples of Biographies

1. James Boswell,  The Life of Samuel Johnson

The biography that ushered in the modern era of true-life writing,  The Life of Samuel Johnson  covered the entirety of its subject’s life, from his birth to his status as England’s preeminent writer to his death. Boswell was a personal acquaintance of Johnson, so he was able to draw on voluminous amounts of personal conversations the two shared.

What also sets this biography apart is, because Boswell was a contemporary of Johnson, readers see Johnson in the context of his own time. He wasn’t some fabled figure that a biographer was writing about centuries later; he was someone to whom the author had access, and Boswell could see the real-world influence his subject had on life in the here and now.

2. Sylvia Nasar,  A Beautiful Mind

Nasar’s 1998 Pulitzer Prize-nominated biography of mathematician John Nash introduced legions of readers to Nash’s remarkable life and genius. The book opens with Nash’s childhood and follows him through his education, career, personal life, and struggles with schizophrenia. It ends with his acceptance of the 1994 Nobel Prize for Economics. In addition to a Pulitzer nomination,  A Beautiful Mind  won the National Book Critics Circle Award for Biography, was a  New York Times  bestseller, and provided the basis for the Academy Award-winning 2001 film of the same name.

3. Catherine Clinton,  Harriet Tubman: The Road to Freedom

Clinton’s biography of the abolitionist icon is a large-scale epic that chronicles Tubman’s singular life. It starts at her birth in the 1820s as the slave Araminta Ross, continuing through her journey to freedom; her pivotal role in the Underground Railroad; her Moses-like persona; and her death in 1913.

Because Tubman could not read or write, she left behind no letters, diaries, or other personal papers in her own hand and voice. Clinton reconstructed Tubman’s history entirely through other source material, and historians often cite this work as the quintessential biography of Tubman’s life.

4. Megan Mayhew Bergman,  Almost Famous Women

Almost Famous Women  is not a biography in the strictest sense of the word; it is a fictional interpretation of real-life women. Each short story revolves around a woman from history with close ties to fame, such as movie star Marlene Dietrich, Standard Oil heiress Marion “Joe” Carstairs, aviatrix Beryl Markham, Oscar Wilde’s niece Dolly, and Lord Byron’s daughter Allegra. Mayhew Bergman imagines these colorful women in equally colorful episodes that put them in a new light—a light that perhaps offers them the honor and homage that history denied them.

Further Resources on Biography

Newsweek  compiled their picks for the  75 Best Biographies of All Time .

The Open Education Database has a list of  75 Biographies to Read Before You Die .

Goodreads put together a list of readers’  best biography selections .

If you’re looking to write biographies,  Infoplease  has instructions for writing shorter pieces, while  The Writer   has practical advice for writing manuscript-length bios.

Ranker  collected  a comprehensive list of famous biographers .

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Definition of biography noun from the Oxford Advanced Learner's Dictionary

• Boswell’s biography of Johnson
• a biography by Antonia Fraser
• The book gives potted biographies of all the major painters.
• blockbuster
• unauthorized
• biography by
• biography of

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Biography: A Very Short Introduction

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Biography: A Very Short Introduction looks at the origins and development of biographical writing. Why do certain people and historical events arouse so much interest? How can biographies be compared with history and works of fiction? Does a biography need to be true? Is it acceptable to omit or conceal things? Does the biographer need to personally know the subject? Must a biographer be subjective? This VSI considers the cultural and historical background of different types of biographies, looking at the factors that affect biographers and whether there are different strategies, ethics, and principles required for writing about one person compared to another.

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Finding Biographical Information

Biographical dictionaries.

• Research Approaches

Biographical Dictionaries by Occupation

Biographical dictionaries by gender, biographical dictionaries by ethnicity, finding more biographical dictionaries.

• Book-length Biographies
• Periodical Articles/Obituaries
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• College/University Alumni/ae Biography
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By no means all biographical dictionaries are covered by the resources listed in Resources of First Resort. More are listed on this page.

• Biograpical Dictionaries by Occupation

Authors including Literary Authors

Contemporary Authors  includes over 116,000 living (or dying since early 1960s) literary and non-literary writers worldwide. Articles typically include short critical essay;  brief personal information; list of author’s writings; critical writings about the author.  New entries appear in  Contemporary Authors  (CA); revisions of CA entries appear in  Contemporary Authors New Revision Series  and the  Permanent Series . HOLLIS Records In  Gale In Context: Biography  (Available to Massachusetts residents via the State Library of Massachusetts)

Dictionary of Literary Biography is a huge series of volumes offering biographical-critical essays on authors, including references for interviews, critical writings. Coverage is international, with emphasis on English and American. Lamont REF.ROOM PN451.D5 1978x (Lamont holds selected volumes.) Widener RR 3129 (Widener holds the complete set.) The DLB is indexed by name in the Literary Index Includes numerous volumes on essayists, travel writers, literary biographers, book collectors and bibliographers, major publishing forms, etc. as well as on literary authors.

American men of science; a biographical directory , 1906- . [title varies: American men & women of science, 1971- ].  HOLLIS Records Internet Archive Full Text  (1905-1970) Internet Archive Full Text   (1971-2009)  Open Year on left to select your year)

Index to editions 1/14: Cabot Science Q141 .A471 1983 CUM. Index 1-14 Widener WID-LC Q141 .A474 (At beginning) Widener | S 120.15.5 (shelved before 14th ed.) Internet Archive Full Text Indexed by  Biography and Genealogy Master Index  (BGMI) from 12th ed. (1971/73) to present.

Complete Dictionary of scientific biography  is the online version of the  Dictionary of scientific biography  and  New Dictionary of Scientific Biography . Offers biographical articles on deceased scientists, including lists of primary and secondary literature. Volume 16 includes a general index and a list of scientists by field. Volumes 17 & 18 are supplements containing scientists left out of the main series and those dying between 1970 and 1981.  New Dictionary of Scientific Biography  contains new articles about scientists dying after 1950 and earlier figures omitted from the original  Dictionary  together with articles updating entries in the original. Unlike the original, psychology and anthropology are covered, with some sociology and economics. --For more recent sources use  History of Science, Technology and Medicine . For new primary works (new editions, etc.) put your person in the author field. For new secondary works, put your person in the Subject field. --Scribner, C. 1980. Publishing the Dictionary of Scientific Biography. Proceedings of the American Philosophical Society, v. 124, no. 5, pp. 320-322.

The following biographical dictionaries either are not included in BGMI or WBIS, or have features (occupational or other indexes) not accessible through these sources.

FemBio . Institute for Women's Biography Research Hannover/Boston, Luise F. Pusch Hannover, Germany --Some entries give substantial information; others are skeletal. Emphasis on European, especially German women. Full entries have source lists, web links, and often portraits and other photographs.

International who's who of women . London: Europa Publications Ltd., 1992- LOCATION: Widener: RR1505.30

Lesser-known women: a biographical dictionary , 1992. LOCATION: Widener: RR4761.21.5 --Includes over 800 living and deceased women arranged chronologically (1600-1991) by year of most significant achievement. Entries are keyed to about 600 numbered sources in a terminal bibliography. Indexes list biographees by name, country, and profession.

Notable American women . Cambridge, MA: Belknap Press of Harvard University Press, 1971-2004. 5 v. LOCATION: Widener: RR 1711. 34 --Covers women deceased before 1999. Articles include extensive source lists giving particular attention to unpublished sources. Classified index (by field) in each vol. Vols. 1-4 indexed in Biography and genealogy master index. Full text in  Women and Social Movements in the United States: 1600-2000

Oxford encyclopedia of women in world history , ed. by Bonnie G. Smith. Oxford; NY: Oxford University Press, 2008 4 v. LOCATION: Widener | RR 1701.60 --Both topical and biographical articles.

Palgrave Macmillan dictionary of women's biography . 4th ed. NY: Palgrave Macmillan, 2005. LOCATION: WID-LC CT3202 .P26 2005 --Includes over 2100 women, living and deceased, from Antiquity to the present. Introductory nine-page survey of other biographical sources. A few entries are dropped with each new edition. Subject index by occupation or reason for prominence. 1st ed.: The International dictionary of women's biography, 1982.

Women in world history: a biographical encyclopedia , ed. by Anne Commire and Deborah Klezmer. Waterford, CT: Yorkin Publications, 1999-2002. 17 v. LOCATION: Widener | RR 1701.50 --Covers living and deceased women, from Antiquity to the present. Most articles have source lists, which include the location of personal papers. The last volume (v. 17) contains indexes by name with variant spellings, by occupation/experience, etc. Indexed in Biography and genealogy master index.

European immigrant women in the United States: a biographical dictionary , ed. by Judy Barrett Litoff, Judith McDonnell. New York : Garland Pub., 1994, 357 p. LOCATION: Widener | WID-LC | E184.E95 L58 1994 --Includes 239 biographies of deceased women who immigrated since the American Revolution. Each biography has a substantial source list, often giving archival sources. Biographee list by area of activity and a short name/topic index.

Jewish women in America: an historical encyclopedia , ed. by Paula E. Hyman and Deborah Dash Moore. NY: Routledge, 1997. 2 v. LOCATION: Widener | WID-LC | DS115.2 .J49 1997x --Long name/topic index and list of biographees by area of activity. Full text in World Biographical Information System.

Latinas in the United States: a historical encyclopedia , ed. by Vicki L. Ruiz and Virginia Sánchez Korrol. Bloomington: Indiana University Press, 2006. 3 v. LOCATION: Widener | WID-LC | E184.S75 L35 2006

Notable Black American women . Detroit: Gale Research, 1992-1996. LOCATION: Widener: RR4781.41.69 Library has: 2 v. --Includes around 1100 women (about half living) with birth dates 1686-1970. Each entry has a source list (which includes archival sources) and many have portraits. There is an "Area of endeavor" list and a detailed subject index in each volume. Indexed in Biography and genealogy master index and full text in Biography in Context

Biographical Indexes

Women in particular: an index to American women , by Kali Herman. Phoenix, AZ: Oryx Press, 1984. LOCATION: Widener: RR 1501. 26 --Index to biographical articles on women in some 54 works of collective biography. Comprises five indexes: (1) Field and career index; (2) Religious affiliation index; (3) Ethnic and racial index; (4) Geographical index; and (5) Alphabetical index. Biographees are entered in as many of the first four sections (and subsections) as are applicable, with full information (name, dates, occupation or field of activity, places of residence, religion, ethnicity, and references to biographical sources) repeated at each entry. The alphabetical (biographee) index lists all the entries in the other four indexes.

Subject guide to women of the world , by Katharine Joan Phenix. Lanham, MD: Scarecrow Press, 1996, 516 p. Location : Widener RR 1501.24.5 --Provides topical and geographical indexes to  Index to women of the world from ancient to modern times; biographies and portraits  (Widener: RR 1501) which provides references to biographical sketches which appear in 945 collective biographies and which is itself indexed by name in Biography and Genealogy Master Index.

African American national biography , ed. by Henry Louis Gates, Jr., Evelyn Brooks-Higginbotham. NY: Oxford University Press, 2013, 12 v. Andover-Harv. Theol | Ref. | E185.96 .A4466 2008 (1st ed.) Lamont | REFERENCE | E185.96 .A4466 2008 (1st ed.) Widener RR 4781.60 (2nd ed.; 2013) Online version  (Continuously updated)

Contemporary Black biography . Vol. 1- Detroit: Gale Research Inc., 1992- LOCATION: Widener: RR 4781.41.59 Substantial articles (two to six pages), with source references. Each volume has about 70 articles. Many entries have photographs or interviews. Covers primarily (but not exclusively) living persons; about 80% are Americans. Indexed by nationality, occupation, subject, and name. Indexed in Biography and genealogy master index

African-American Biographical Database  is an online collection of about 300 full-text biographical dictionaries. Also available as a microfiche set, with printed index, in Widener's Reference Room (as  Black Biographical Dictionaries 1790-1950 ).

Notable Asian Americans , ed by Helen Zia and Susan B. Gall. New York : Gale Research, 1995, 468 p. HOLLIS Record

Distinguished Asian Americans: a biographical dictionary , ed. by Hyung-chan Kim. Westport, CT: Greenwood Press, 1999, 430 p. HOLLIS Record

Who's who among Asian Americans . Detroit : Gale Research Inc., 1994. 1 v.  HOLLIS Record

Asian and Pacific Islander Americans , ed. by Gary Y. Okihiro. Ispwich, Mass.: Salem Press, 2013. 3 v. HOLLIS Record

Native American biographies  (North American Indian Thought and Culture) 

Find unindexed works of collective biography in your subject area by searching the  HOLLIS Catalog  and  WorldCat , a collective catalog of thousands of libraries. Very specialized and very recent collective biographies are least likely to be indexed.

To find biographical dictionaries and who's whos, attach the words Biography AND (Dictionaries OR Encyclopedias OR "bio-bibliographies") to your search term, thus:

Physicians AND Biography AND (Dictionaries OR Encyclopedias OR "Bio-bibliography")

Some collective biographical works, especially those including a small number of persons,  carry the term Biography only.  These can only be found by browsing through the whole (and large)  Physicians Biography set

Records for who's whos sometimes, but by no means consistently, bear the subdivisions Directories or Periodicals.

When you have found a likely collective biography, check the lists of indexed sources in  Biography and Genealogy Master Index  (Sources Indexed) and do a Bibliographical search in the  World Biographical Information System (WBIS) . Although many biographical dictionaries are available full text in WBIS, the print versions often contain vocational/geographical indexes and other features not available through WBIS.

Subject dictionaries and encyclopedias, although not purely biographical, often include biographical articles

Example: History of physical anthropology : an encyclopedia / edited by Frank Spencer. Subject: Physical anthropology Encyclopedias.

Search for them in HOLLIS+: anthropology AND (encyclopedias OR dictionaries)

Bibliographies of Collective Biography

The following sources list biographical dictionaries by subject:

Biographical dictionaries and related works: an international bibliography of more than 16,000 collective biographies  ..., by Robert B. Slocum, 2nd ed. Detroit, Mich.: Gale Research Co., 1986. LOCATION: Law School: Special Coll Ref CT 104.S55x 1986 LOCATION: Widener: RR 1501. 13.2 Library has: 2 v. --Thorough and conveniently arranged, by geography and profession, bibliography of pre-1986 biographical dictionaries.

Guide to reference in genealogy and biography , ed. by Mary K. Mannix, Fred Burchsted, and Jo Bell Whitlatch. Chicago: ALA Editions, an imprint of the American Library Association, 2015, 372 pages. HOLLIS Record Biographical section arranged by continent and country.

ARBA guide to biographical dictionaries . Littleton, Colo.: Libraries Unlimited, 1986. LOCATION: Widener: WID-LC CT103.Z99 A73 x, 1986 --Continued by ARBA guide to biographical resources, 1986-1997 . Englewood, Colo.: Libraries Unlimited, 1998. LOCATION: Widener: RR 1501.13.6

Bibliographie analytique des biographies collectives imprimées de la France contemporaine, 1789-1985 , by Alfred Fierro. Geneve: Slatkine, 1986. LOCATION: Andover-Harv. Theol: Ref. Z2170.F45 1986 LOCATION: Widener: RR 1771.30 --Sources arranged by profession.

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Biological Information

Since the 1950s, the concept of information has acquired a strikingly prominent role in many parts of biology. This enthusiasm extends far beyond domains where the concept might seem to have an obvious application, such as the biological study of perception, cognition, and language, and now reaches into the most basic parts of biological theory. Hormones and other cellular products through which physiological systems are regulated are typically described as signals. Descriptions of how genes play their causal role in metabolic processes and development are routinely given in terms of “transcription”, “translation”, and “editing”. The most general term used for the processes by which genes exert their effects is “gene expression”. The fates of cells in a developing organism are explained in terms of their processing of “positional information” given to them from surrounding cells and other factors. Many biologists think of the developmental processes by which organisms progress from egg to adult in terms of the execution of a “developmental program”. Other biologists have argued for a pivotal role for information in evolution rather than development: John Maynard Smith and Eors Szathmáry (for example) suggest that major transitions in evolution depend on expansions in the amount and accuracy with which information is transmitted across generations. And some have argued that we can only understand the evolutionary role of genes by recognizing an informational “domain” that exists alongside the domain of matter and energy.

Both philosophers and biologists have contributed to an ongoing foundational discussion of the status of this mode of description in biology. It is generally agreed that the sense of information isolated by Claude Shannon and used in mathematical information theory is legitimate, useful, and relevant in many parts of biology. In this sense, anything is a source of information if it has a range of possible states, and one variable carries information about another to the extent that their states are physically correlated. But it is also agreed that many uses of informational language in biology seem to make use of a richer and more problematic concept than Shannon’s. Some have drawn on the teleosemantic tradition in philosophy of mind to make sense of this richer concept. Other theorists have countered that Shannon’s correlational conception of information is richer than it looks.

A minority tradition has argued that the enthusiasm for information in biology has been a serious theoretical wrong turn, and that it fosters naive genetic determinism, other distortions of our understanding of the roles of interacting causes, or an implicitly dualist ontology. However, this sceptical response is fading, with key sceptics coming to accept a modest but genuine role for informational concepts in the life sciences. Others have taken the critique seriously but tried to distinguish legitimate appeals to information from misleading or erroneous ones.

1. Introduction

2. shannon’s concept of information, 3. teleosemantic and other richer concepts, 4. the genetic code, 5. signalling systems, 6. rejections of informational concepts in biology, 7. genetic programs, 8. information and evolution, other internet resources, related entries.

Biology is concerned with living organisms—with their structure, activities, distribution in space and time, and participation in evolutionary and developmental histories. Many of these organisms engage in activities that seem best understood in terms of information processing or representation. These include perception, cognition, signalling, and language use. We will not much be concerned with the role of concepts of information and representation in these cognitive contexts. For over the second half of the 20 th century, biology came to apply informational concepts (and their relatives) far more broadly than this. For many biologists, the most basic processes characteristic of living organisms should now be understood in terms of the expression of information: the response to signals, the execution of programs, and the interpretation of codes. So although contemporary mainstream biology is an overtly materialist field, it has come to employ concepts that seem intentional or semantic. These come with a long history of causing foundational problems for materialists (and, to some extent, for everyone else).

The embrace of informational and other semantic concepts has been especially marked within genetics, and other fields—evolutionary theory and developmental biology—with strong ties to genetics. But informational language is not only found there: for example, hormones are routinely seen as long distance signals that enable one organ system to coordinate with others (Levy 2011). That said, the usage that has generated the most discussion is found in the description of the relations between genes and the various structures and processes to which genes contribute. For many biologists, the causal role of genes should be understood in terms of their carrying information about their various products; and perhaps as well about the environments in which these products enhance fitness (Lorenz 1965; Shea 2013). The expression of that information might depend on the presence of various environmental factors, but the same can be said of other kinds of message. Breaking things down further, we can recognize two causal roles for genes, and hence two potential explanatory roles for genetic information, within biology. Genes are crucial to both explaining the development of individual organisms, and to explaining the inheritance of characteristics across generations. Information has been invoked in both explanatory contexts.

One important use of informational language is relatively uncontroversial, through being anchored in striking and well established facts about the role of DNA and RNA in the manufacture of protein molecules in cells, a set of facts summarized in the familiar chart representing the “genetic code”, that maps triplets of the DNA bases (C, A, T, G) to individual amino acids, which are the building blocks of protein molecules. Not even this use of “coding” language is completely uncontroversial (Sarkar 1996). More importantly, enthusiasm for the use of informational language in biology both predates the discovery of the DNA-RNA-amino acid mapping (Schrödinger 1944), and goes far beyond it, so this mapping is only a partial explanation of the uptake of informational notions in biology. Current applications of informational concepts in biology include:

• The description of whole-organism phenotypic traits (including complex behavioral traits) as specified or coded by information contained in the genes,
• The treatment of many causal processes within cells, and perhaps of the whole-organism developmental sequence, in terms of the execution of a program stored in the genes,
• Treating the transmission of genes (and sometimes other inherited structures) as a flow of information from the parental generation to the offspring generation.
• The idea that genes themselves, for the purpose of evolutionary theorizing, should be seen as, in some sense, “made” of information. Information becomes a fundamental ingredient in the biological world.
• Characterising, in a fully general way, the dynamics of idealized populations changing as a result of natural selection.

There is no consensus about the status of these ideas. Indeed, the use of informational notions is controversial even when giving accounts of animal communication, with some theorists denying that such communication is the flow of information from one animal to another (Krebs and Dawkins 1984; Owren et al. 2010). The result has been a growing foundational discussion within biology and the philosophy of biology. Some have hailed the employment of informational concepts as a crucial advance (Williams 1992). Others have seen almost every biological application of informational concepts as a serious error, one that distorts our understanding and contributes to lingering genetic determinism (Francis 2003). Most of the possible options between these extreme views have also been defended. Perhaps most commentators within philosophy have seen their project as one of sorting through the various kinds of informational description that have become current, distinguishing legitimate ones from illegitimate ones (Godfrey-Smith 2000; Griffiths 2001; Godfrey-Smith 2007; Shea 2013; Lean 2014). Philosophers have also tried to give a reductive or naturalistic explanation for the legitimate ones.

This entry proceeds as follows. In the next section we discuss the most unproblematic technical use of information in biology, which draws on Shannon and the mathematical theory of information. Against that background, some of the more contentious uses are both motivated and introduced. In section 3 , we discuss recent attempts to build a richer concept out of Shannon’s technical notion. We then discuss the status of the “genetic code” in its original sense ( section 4 ), and signalling systems inside and outside of genetics ( section 5 ). We then look at ways of rejecting informational forms of description ( section 6 ). The key sceptical idea is the so-called “parity thesis”: the idea that while genes play an immensely important role in evolution and development, so do other factors, and so it is a mistake to think of genes and only genes as instructing or controlling these processes. The final two sections discuss the idea that biological development in individuals proceeds by the execution of a program ( section 7 ), and the use of information in understanding the role of inheritance systems in the “major transitions” in evolution ( section 8 ).

It is common to begin the analysis of information in biology with an uncontroversial but minimal notion: a causal or correlational conception. Smoke in the air and fire in the forest are correlated with one another. If you see a forest burning at night, you can predict the air will be smoky. If you see a plume of smoke rising from above the trees, you can predict that below there is a fire. The correlation does not have to be perfect to be informative. Putting the point generally, a signal carries information about a source, in this sense, if we can predict the state of the source from the signal. This sense of information is associated with Claude Shannon (1948) who showed how the concept of information could be used to quantify facts about contingency and correlation in a useful way, initially for use in communication technology. For Shannon, anything is a source of information if it has a number of alternative states that might be realized on a particular occasion. Any other variable contains some information about that source, or carries information about it, if its state is correlated with the state of the source. This is a matter of degree; a signal carries more information about a source if its state is a better predictor of the source, less information if it is a worse predictor.

In this sense, a signal can carry information about a source without there being any biological system designed to produce that signal, nor any to use it once produced. When a biologist employs this sense of information in a description of gene action or of other processes, he or she is just adopting a quantitative framework for describing ordinary correlations or causal connections within those systems. So it is one thing for there to be a signal carrying information about a source; quite another to explain biological processes as the result of signalling. The familiar example of tree rings is helpful here. When a tree lays down rings, it establishes a structure that can be used by us to make inferences about the past. The number and size of the rings carry information, in Shannon’s sense, about the history of the tree and its circumstances (the technique, “dendrochronology”, is scientifically important). Despite the usefulness of the informational description, there is no sense in which we are explaining how the tree does what it does in informational terms. The only reader or user of the information in the tree rings is the human observer. The tree itself does not exploit the information in its rings to control its growth or flowering. Similarly, we might note ways in which the distribution of different DNA sequences within and between biological populations “carries information about” the historical relationships between those populations, and the histories of the individual populations themselves. Using this information has given evolutionary biologists a much richer and more reliable picture of the history of life (for a good introduction to these uses see Bromham 2008). It has been argued, for example, that the greater diversity in mitochondrial DNA in African populations, in comparison to other human populations, is an indicator of a comparatively recent human dispersal from an African origin. In making these inferences, informational concepts can be useful tools. But this is just a more complicated version of what is going on in the case of tree rings. The appeal to information has an inferential use that is no way explanatory . A large proportion of the informational descriptions found in biology have this character.

The border between these two categories, however, can be hard to discern. Steven Frank (2012) has developed a detailed mapping between equations describing change due to natural selection and equations used in information theory. Change across generations can be seen as the accumulation of information by a population about its environment. So far, there is no claim that populations use this information in any way, and the situation is akin to that found in tree rings: a tree accumulates, in its rings, information about climatic conditions in its past, though the tree does nothing with that information. Frank may view the role of information acquired by evolution in a more substantive way: “selection causes populations to accumulate information about the fit of characters to the environment” (2012: 2391). But it is doubtful that anything more than an inferential role has yet been shown for this kind of information.

Consequently, philosophers have sometimes set the discussion up by saying that there is one kind of “information” appealed to in biology, the kind originally described by Shannon, that is unproblematic and does not require much philosophical attention. The term “causal” information is also sometimes used to refer to this concept, and this is essentially Grice’s “natural meaning”, (see entry on Grice ) from his work in the philosophy of language (Grice 1957). Information in this sense exists whenever there is contingency and correlation. So we can say that genes contain information about the proteins they make, and also that genes contain information about the whole-organism phenotype. But when we say that, we are saying no more than what we are saying when we say that there is an informational connection between smoke and fire, or between tree rings and a tree’s age. The more contentious question becomes whether or not biology needs another , richer concept of information as well. Information in this richer sense is sometimes called “semantic” or “intentional” information.

Why might we think that biology needs to employ a richer concept? One thought is that genes play a special, instructional role in development, telling the embryo how to grow. It is true that genes carry Shannon-information about phenotypes: the genome of a fertilized egg predicts much of the resulting phenotype. In mammals, for example, chromosome structure predicts the sex of the adult animal. But if an informational relationship between gene and phenotype is supposed to involve a distinctive instruction-like mode of causation, then this cannot be information in Shannon’s sense. For environmental factors, not just genes, “carry information” about phenotypes, in Shannon’s sense. With respect to Shannon information, there is a “parity” between the roles of environmental and genetic causes (Griffiths and Gray 1994). Moreover, if we are using Shannon’s concept, then the informational relationship between genotype and phenotype is symmetrical. For example, once a reader knows that both authors of this article are male, they can predict that we both carry a Y-chromosome. Some talk about information in biology is consistent with these features of Shannon information, but some is not. In particular, it is usually thought that at least some applications of informational language to genes ascribe to them a property that is not ascribed to environmental conditions, even when the environment is predictively important.

In addition, a message that carries “semantic information”, it is often thought, has the capacity to mis -represent, as well as accurately represent, what it is about. There is a capacity for error. Shannon information does not have that feature; we cannot say that one variable “carried false information” about another, if we are using Shannon’s sense of the term. But biologists do apparently want to use language of that kind when talking about genes. Genes carry a message that is supposed to be expressed, whether or not it actually is expressed. Thus the developmental disorders caused by thalidomide in the 1960s were due to an inability to use genetic signals that were supposed to control limb development. These genes were present and active; thalidomide did not do its harm by causing mutations.

These are the usual “marks” that are taken in the literature to show that a richer sense of information than Shannon’s has been introduced to biology. But the most crucial difference between the less and more contentious applications of informational concepts is that, in the richer cases, information use is supposed to help explain how biological systems do what they do—how cells work, how an egg can develop into an adult, how genetic inheritance mechanisms make the evolution of complex phenotypes possible.

At this point, there are a number of options on the table. One is to deny that genes, cells and other biological structures literally traffic in information in ways that explain their behaviour, but to argue nevertheless that this is a useful analogy or model. The idea is that there are useful similarities between paradigm cases of information and representation using systems—cognitively sophisticated agents, thinking and communicating with one another—and biological systems. Hormones, for example, are usefully thought of as messages because they are small, stable, energetically inexpensive items that can travel long distances (relative to their own size) without decaying, to specialized locations where they have predictable, specific, and important effects on arrival. But while this is a helpful way of thinking, perhaps we should not take talk of messages too seriously. For example, we should not treat a question about whether prolactin is a report of pregnancy or an instruction to the mammary glands as if it had a literally correct answer. Arnon Levy has developed the most sophisticated version of this view of biological information (Levy 2011).

A second option is to argue that genes and other biological structures literally carry semantic information, and their informational character explains the distinctive role of these structures in biological processes. If we think of genes or cells as literally carrying semantic information, our problem changes; who or what could count as composing or reading these messages? Paradigm cases of structures with semantic information—pictures, sentences, programs—are built by the thought and action of intelligent agents. So we need to show how genes and cells—neither intelligent systems themselves nor the products of intelligence—can carry semantic information, and how the information they carry explains their biological role. We need some kind of reductive explanation of semantic information (arguably, we need this to understand cognition, too). One place we might look for such an analysis is naturalistic philosophy of mind.

A third option is to argue that causal information itself can explain biological phenomena, and no additional concept of information is needed. Biological systems, on this view, can be adapted to send or receive signals that carry causal information. Elevated prolactin, for example, does covary with pregnancy, and that is no accident. Likewise, the production of milk is a designed response to the registration of these elevated levels at the mammary glands. While it is true that lactation carries as much causal information about prolactin levels as those levels carry about milk flow, there is a physical asymmetry, hence directionality, between source and receiver. (After birth, suckling stimulates local production of prolactin, so the physical asymmetry is the signal of pregnancy from the pituitary gland to the breasts, tuning them for lactation). Brian Skyrms’ work has been very important in returning causal information to centre stage, in part because his work shows that sender-receiver signalling systems need not be cognitively sophisticated agents (Skyrms 2010). Sending and receiving causal information can emerge and stabilise amongst simple systems; certainly amongst systems no more complex than a cell.

As noted above, several philosophers and biologists have argued that much informational talk about genes uses a richer concept than Shannon’s, but this concept can be given a naturalistic analysis. The aim has been to make sense of the idea that genes semantically specify their normal products, in a sense similar to that seen in some paradigm cases of symbolic phenomena.

If genes are seen as “carrying a message” in this sense, the message apparently has a prescriptive or imperative content, as opposed to a descriptive or indicative one. Their “direction of fit” to their effects is such that if the genes and the eventual structure produced (the phenotype) do not match, what we have is a case of unfulfilled instructions rather than inaccurate descriptions. Alternatively, perhaps it is possible to think that the genes are telling the developing phenotype the environment it can expect. For the gene pool from which those genes have been drawn has been filtered by selection. In arid regions of Australia, genes which contributed to the development of leaves whose shape and surface restricted water loss have become common. Hence perhaps we can see these genes as telling the trees: conditions will be dry (for this line of thought, see Shea 2011, 2013).

In making sense of these ideas, the usual way to proceed has been to make use of a rich concept of biological function , in which the function of an entity derives from a history of natural selection (see Teleological Notions in Biology ). This move is familiar from the philosophy of mind, where similar problems arise in the explanation of the semantic properties of mental states (Millikan 1984). When an entity has been subject to and shaped by a history of natural selection, this can provide the grounding for a kind of purposive or normative description of the causal capacities of that entity. To use the standard example (Wright 1973), the function of a heart is to pump blood, not to make thumping sounds, because it is the former effect that has led to hearts being favored by natural selection. The hope is that a similar “teleofunctional” strategy might help make sense of the semantic properties of genes, and perhaps other biological structures with semantic properties.

The idea of a teleosemantic approach to genetic information was developed in an early form by Sterelny et al. (1996); see also Maclaurin (1998). The eminent biologist John Maynard Smith took a similar approach, when he tried to make sense of his own enthusiasm for informational concepts in biology (Maynard Smith 2000; and see also the commentaries that follow Maynard Smith’s article). Eva Jablonka has also defended a version of this idea (Jablonka 2002). Her treatment is more unorthodox, as she seeks to treat environmental signals as having semantic information, along with genes, if they are used by the organism in an appropriate way. Nick Shea has developed the most sophisticated teleosemantic treatment of genes to date (see Shea 2007a,b, 2011, 2013).

One way of developing these ideas is to focus on the evolved functions of the genetic machinery as a whole (Godfrey-Smith 1999). Carl Bergstrom and Martin Rosvall take this approach, emphasising the adapted, impressively engineered features of intergenerational gene flow in developing their “transmission sense of information”. Bergstron and Rosvall point out that intergeneration gene flow is structured so as to make possible the transmission of arbitrary sequences (so the message is relatively unconstrained by the medium); that information is stored compactly and stably; that the bandwidth is large and indefinitely extendable. DNA sequences are reliably replicable with very high fidelity, so transmission is accurate, yet the mapping between DNA and amino acid sequence seems optimised to reduce the impact of those errors that do occur. Many point mutations are mapped onto the same or a chemically similar amino acid. Bergstrom and Rosvall conclude both that DNA replication is an exquisitely designed information channel, and that we can tell this from the characteristics of the DNA-amino acid system itself. We do not need to know what signals from the parental generation say to the offspring generation, in order to know that the characteristics of DNA replication are explained by its information-carrying capacities (Bergstrom and Rosvall 2011).

The more usual route, and the one taken both by Sterelny et al. (1996) and by Maynard Smith (2000) is to focus on the natural selection of particular genetic elements. That view faces an immediate problem, for the fact that specific genetic elements (or the genetic system as a whole) have an evolved function is clearly not sufficient for genes to carry semantic information. Legs are for walking, but they do not represent walking. Enzymes are for catalyzing reactions, but they do not instruct this activity. There are things that legs and enzymes are supposed to do, but this does not make them into information-carriers, in a rich beyond-Shannon sense. Why should it do so for genes? Sterelny, Smith and Dickison (1996), suggested that the differences between genes and legs is that genes have been selected to play a causal role in developmental processes. They add, however, that any non-genetic factors that have a similar developmental role, and have been selected to play that role, also have semantic properties. So Sterelny, Smith and Dickison want to ascribe very rich semantic properties to genes, but not only to genes. Some non-genetic factors have the same status. Even so, many quite plausible cases turn out not to be informational on this view: prolactin, like most hormones, does not have a specifically developmental function, so it would not count as carrying information to the mammary glands. This is one reason why Levy thinks of talk of information as merely metaphorical (Levy 2011). In contrast to the views of Sterelny, Smith and Dickison, in his 2000 paper, John Maynard Smith argued that only genes carry semantic information about phenotypes. He suggested that in contrast to other developmental resources, the relationship between adapted gene and phenotypic outcome is arbitrary; the gene-trait relationship is like the word-object relationship. This idea is intriguing but hard to make precise. One problem is that any causal relation can look “arbitrary” if it operates via many intervening links, for there are many possible interventions on those links which would change the product of the causal chain. The problem of arbitrariness is discussed further in section 4 .

To distinguish between ordinary biological functions and representational functions, Nicholas Shea draws upon the more elaborate machinery of Ruth Millikan’s teleosemantic theory. For Millikan, any object that has semantic properties plays a role that involves a kind of mediation between two “cooperating devices”, a producer and a consumer device. In the case of an indicative signal or representation, the representation is supposed to affect the activities of the consumer in a way that will only further the performance of the consumer’s biological functions if some state of affairs obtains. In the case of an imperative representation, the representation is supposed to affect the activities of the consumer by inducing them to bring some state of affairs about. Shea treats genetic messages as having both indicative and imperative content, and depending on both producers and consumers (Shea 2007b, 2011, 2013). Where Shea follows Millikan in emphasizing the roles of both producers and consumers, Jablonka (2002) tries to achieve as much as possible with an emphasis on consumer mechanisms alone. It seems clear that the attention to producer and consumer mechanisms is a step forward in this discussion, but we will see in section 5 that it also poses problems. In thinking about both inheritance and development, it turns out to be unclear whether there are independently identifiable mechanisms which count as senders and receivers, producers and consumers.

The overall picture envisaged by the teleosemantic approach has undeniably appealing structural features. If this program succeeds, we would have an uncontroversial sense of information, via Shannon, that applies to all sorts of physical correlations. This picture can then be developed by identifying a subset of cases in which these signals have been co-opted or produced to drive biological processes. In addition, perhaps we can appeal to rich semantic properties in cases where we have the right kind of history of natural selection to explain the distinctive role of genes, and perhaps other factors, in development. Genes and a handful of non-genetic factors would have these properties; most environmental features that have a causal role in biological development would not. There remain many problems of detail, but the appeal of the overall picture provides, at least for some, good reason to persevere with some account along these lines.

So far we have mostly discussed the concept of information; there has not been much talk of “coding”. With the exception of our discussion of Bergstrom and Rosvall’s “transmission sense” of information, the discussion so far has not emphasised the distinctive features of the cell-level processes in which genes figure, such as the language-like combinatorial structure of the “genetic code”. These structural features of DNA and its relation to amino acids are not central to some of the ideas about information in biology, even when the focus is on development and inheritance. As noted above, the enthusiasm for semantic characterization of biological structures extends back before the genetic code was discovered (see Kay 2000 for a detailed historical treatment). But one line of thought in the literature, overlapping with the ideas above, has focused on the special features of genetic mechanisms, and on the idea of “genetic coding” as a contingent feature of these mechanisms.

Both Peter Godfrey-Smith and Paul Griffiths have argued that there is one highly restricted use of a fairly rich semantic notion within genetics that is justified (Godfrey-Smith 2000; Griffiths 2001). This is the idea that genes “code for” the amino acid sequence of protein molecules, in virtue of the peculiar features of the “transcription and translation” mechanisms found within cells. Genes specify amino acid sequence via a templating process that involves a regular mapping rule between two quite different kinds of molecules (nucleic acid bases and amino acids). This mapping rule is combinatorial , and apparently arbitrary (in a sense that is hard to make precise, though see Stegmann 2004 for discussion of different versions of this idea).

Figure 1 below has the standard genetic code summarized. The three-letter abbreviations such as “Phe” and “Leu” are types of amino acid molecules.

Figure 1. The Standard Genetic Code

This very narrow understanding of the informational properties of genes is basically in accordance with the influential early proposal of Francis Crick (1958). The argument is that these low-level mechanistic features make gene expression into a causal process that has significant analogies to paradigmatic symbolic phenomena.

Some have argued that this analogy becomes questionable once we move from the genetics of simple prokaryotic organisms (bacteria and archaea), to those in eukaryotic cells (Sarkar 1996). Mainstream biology tends to regard the complications that arise in the case of eukaryotes as mere details that do not compromise the basic picture we have of how gene expression works (for an extensive discussion of these complexities, by those who think they really matter, see (Griffiths and Stotz 2013)). An example is the editing and “splicing” of mRNA transcripts. The initial stage in gene expression is the use of DNA in a template process to construct an intermediate molecule, mRNA or “messenger RNA”, that is then used as a template in the manufacture of a protein. The protein is made by stringing a number of amino acid molecules together. In eukaryotes the mRNA is often extensively modified (“edited”) prior to its use. Moreover, much of the DNA in a eukaryotic organism is not transcribed and translated at all. Some of this “non-coding” DNA (note the informational language again) is certainly functional, serving as binding sites for proteins that bind to the DNA thus regulating the timing and rate of transcription. The extent of wholly nonfunctional DNA remains unclear. These facts make eukaryotic DNA a much less straightforward predictor of the protein’s amino acid sequence than it is in bacteria, but it can be argued that this does not much affect the crucial features of gene expression mechanisms that motivate the introduction of a symbolic or semantic mode of description.

So the argument in Godfrey-Smith (2000) and Griffiths (2001) is that there is one kind of informational or semantic property that genes and only genes have: coding for the amino acid sequences of protein molecules. But this relation “reaches” only as far as the amino acid sequence. It does not vindicate the idea that genes code for whole-organism phenotypes, let alone provide a basis for the wholesale use of informational or semantic language in biology. Genes can have a reliable causal role in the production of a whole-organism phenotype, of course. But if this causal relation is to be described in informational terms, then it is a matter of ordinary Shannon information, which applies to environmental factors as well. That said, it is possible to argue that the specificity of gene action, and the existence of an array of actual and possible alternatives at a given site on a chromosome, means that genes exert fine-grained causal control over phenotypes, and that few other developmental resources exert this form of causal control (Waters 2007; Maclaurin 1998; Stegmann 2014). In contrast, Griffiths and Stotz (2013) say that these other factors are often a source of “Crick information”, as they contribute to specifying the linear sequence of a gene product. We return to this issue in section 6 .

One of the most appealing, but potentially problematic, features of the idea that genes code for amino acid sequences concerns the alleged “arbitrariness” of the genetic code. The notion of arbitrariness figures in other discussions of genetic information as well (Maynard Smith 2000; Sarkar 2003; Stegmann 2004). It is common to say that the standard genetic code has arbitrary features, as many other mappings between DNA base triplets and amino acids would be biologically possible, if there were compensating changes in the machinery by which “translation” of the genetic message is achieved. Francis Crick suggested that the structure of the genetic code should be seen as a “frozen accident”, one that was initially highly contingent but is now very hard for evolution to change (Crick 1958). But the very idea of arbitrariness, and the hypothesis of a frozen accident, have become problematic. For one thing, as noted in our discussion of Bergstrom and Rosvall (2011) above, the code is not arbitrary in the sense that many others would work as well. To the contrary, the existing code is near-optimal for minimising error costs. Conceptually, it seems that any causal relation can look “arbitrary” if it operates via many intervening links. There is nothing “arbitrary” about the mechanisms by which each molecular binding event occurs. What makes the code seem arbitrary is the fact that the mapping between base triplets and amino acids is mediated by a causal chain with a number of intervening links (especially involving “transfer RNA” molecules, and enzymes that bind amino acids to these intervening molecules). Because we often focus on the “long-distance” connection between DNA and protein, the causal relation appears arbitrary. If we focused on steps in any other biological cascade that are separated by three or four intervening links, the causal relation would look just as “arbitrary”. So the very idea of arbitrariness is elusive. And empirically, the standard genetic code is turning out to have more systematic and non-accidental structure than people had once supposed (Knight, Freeland, and Landweber 1999). The notion of arbitrariness has also been used in discussions of the links between genes and phenotypes in a more general sense. Kirscher and Gerhart (2005) discuss a kind of arbitrariness that derives from the details of protein molecules and their relation to gene regulation. Proteins that regulate gene action tend to have distinct binding sites, which evolution can change independently. To bind, a protein must be able to attach to a site, but that requires congruence only with a local feature of the protein, not sharply constraining its overall shape (Planer 2014). This gives rise to a huge range of possible processes of gene regulation. So there is a perennial temptation to appeal to the idea of arbitrariness when discussing the alleged informational nature of some biological causation.

In section 2 , we noted that Brian Skyrms’ work on signalling systems has made this framework a very natural fit for a quite large range of biological phenomena. As we remarked above, it is very intuitive to see hormones such as insulin, testosterone, and growth hormone as signals, as they are produced in one part of the body, and travel to other parts where they interact with “receptors” in a way that modifies the activities of various other structures. It is routine to describe hormones as “chemical messages”. The Skyrms framework fits these designed cause and response systems within biological agents for three reasons. First, as noted, this framework shows that signalling does not require intelligence or an intelligent grasp of signal meaning.

Second, the simplest cases for models of signalling are cases in which there is common interest. The sender and receiver are advantaged or disadvantaged by the same outcomes. While complete common interest is atypical of organism-to-organism communication, the cells and other structures within an organism share a common fate (with complex exceptions). So in this respect the base models might fit organ-to-organ communication better than they fit between-organism phenomena.

Third, in many of these biological systems, the abstract structure specified as a signalling system—environmental source, sender, message, receiver, response—maps very naturally onto concrete biological mechanisms. For example, Ron Planer has recently argued that we should see gene expression as the operation of a signalling system (Planer 2014). His view is quite nuanced, for the identity of the sender, receiver, and message vary somewhat from case to case. For example, when a protein is a transcription factor, then the gene counts as a sender. He treats other gene-protein relations differently. The details of his view need not concern us here. The point is that there is machinery in the cell—genes, proteins mRNA transcripts, ribosomes and their associated tRNA—that can be plausibly mapped onto sender-receiver systems. There is nothing forced about mapping the information-processing sender-receiver structure onto the molecular machinery of the cell.

However, while this framework very naturally fits within cell and between cell processes, it is much less clear how naturally other suggestions mesh with this framework. For example, in the Bergstrom-Rosvall picture of intergenerational transmission, who or what are the senders and receivers? Perhaps in the case of multicelled organisms, the receiver exists independently of and prior to the message. For an egg is a complex and highly structured system, before gene expression begins in the fertilised nucleus, and that structure plays an important role in guiding that gene expression (Sterelny 2009; Gilbert 2013). But most organisms are single celled prokaryotes, and when they fission, it is not obvious that there is a daughter who exists prior to and independently of the intergenerational genetic message she is to receive.

Likewise, Nicholas Shea’s Millikan-derived analysis does not seem to map naturally onto independently specifiable biological mechanisms. For him, the sender of genetic messages is natural selection operating on and filtering the gene pool of a population; messages are read by the developmental system of the organism as a whole (Shea 2013). But the less clearly a sender-receiver or producer-consumer framework maps onto independently recognised biological mechanisms, the more plausible a fictionalist or analogical analysis of that case becomes. So we see an important difference between a reader being the developmental system as a whole, and reader being a receptor on a cell membrane. The cell-level machinery of transcription and translation (the ribosomal/tRNA machinery, especially) really is a reader or consumer of nucleic acid sequences, with the function of creating protein products that will have a variety of uses elsewhere in the cell. But this realization of the causal schematism applies only at the cell level, at the level at which the transcription and translation apparatus shows up as a definite part of the machinery. One of the most extraordinary features of ontogeny is that it proceeds reliably and predictably without any central control of the development of the organism as a whole. There is nothing, for example, that checks whether the left-side limbs are the same size as the right side limbs, intervening to ensure symmetry. Similarly, there are DNA sequence-readers, and some intercellular sender-receiver systems with clear “readers” (such as neural structures), but there are no higher-level readers that interpret a message specifying the structure of a whole limb-bud in the embryo.

Some biologists and philosophers have argued that the introduction of informational and semantic concepts has had a bad effect on biology, that it fosters various kinds of explanatory illusions and distortions, perhaps along with ontological confusion. Here we will survey some of the more emphatic claims of this kind, but some degree of unease can be detected in many other discussions (see, for example, Griesemer 2005).

The movement known as Developmental Systems Theory (DST) has often opposed the mainstream uses of informational concepts in biology, largely because of the idea that these concepts distort our understanding of the causal processes in which genes are involved. For a seminal discussion, see Oyama (1985), and also Lehrman (1970); Griffiths and Gray (1994); Griffiths and Neumann-Held (1999). These theorists have two connected objections to the biological use of informational notions. One is the idea that informational models are preformationist. Preformationism, in its original form, in effect reduces development to growth: within the fertilized egg there exists a miniature form of the adult to come. Preformationism does not explain how an organized, differentiated adult develops from a much less organized and more homogeneous egg; it denies the phenomenon. DST’s defenders suspect that informational models of development do the same. In supposing, for example, that instructions for a “language organ” are coded in the genome of a new-born baby, you do not explain how linguistic abilities can develop in an organism that lacks them, and you foster the illusion that no such explanation is necessary. (See Francis 2003 for a particularly vigorous version of the idea that the appeal to information leads to pseudo-explanation in biology.)

Second, DST theorists have often endorsed a “parity thesis”: genes play an indispensable role in development, but so do other causal factors, and there is no reason to privilege gene’s contribution to development. This claim is often buttressed by reference to Richard Lewontin’s arguments for the complexity and context sensitivity of developmental interaction, and his consequent arguments that we cannot normally partition the causal responsibility of the genetic and the environmental contributions to specific phenotypic outcomes (Lewontin 1974, 2000). DST theorists think that informational models of genes and gene action make it very tempting to neglect parity, and to attribute a kind of causal primacy to these factors, even though they are just one of a set of essential contributors to the process in question. Once one factor in a complex system is seen in informational terms, the other factors tend to be treated as mere background, as supports rather than bona fide causal actors. It becomes natural to think that the genes direct, control, or organise development; other factors provide essential resources. But, the argument goes, in biological systems the causal role of genes is in fact tightly interconnected with the roles of many other factors (often loosely lumped together as “environmental”). Sometimes a gene will have a reliable effect against a wide range of environmental backgrounds; sometimes an environmental factor will have a reliable effect against a wide range of genetic backgrounds. Sometimes both genetic and environmental causes are highly context-sensitive in their operation. Paul Griffiths has emphasised this issue, arguing that the informational mode of describing genes can foster the appearance of context-independence:

Genes are instructions—they provide information—whilst other causal factors are merely material…. A gay gene is an instruction to be gay even when [because of other factors] the person is straight. (Griffiths 2001: 395–96)

The inferential habits and associations that tend to go along with the use of informational or semantic concepts are claimed to lead us to think of genes as having an additional and subtle form of extra causal specificity. These habits can have an effect even when people are willing to overtly accept context-dependence of (most) causes in complex biological systems. So DST theorists suggest that it is misleading to treat genes and only genes as carrying “messages” that are expressed in their effects. To say this is almost inevitably to treat environmental factors as secondary players.

The parity thesis has been the focus of considerable discussion and response. In a helpful paper, Ulrich Stegmann shows that the parity thesis is really a cluster of theses rather than a single thesis (Stegmann 2012). Some ways of interpreting parity make the idea quite uncontroversial, as no more than an insistence on the complex and interactive character of development, or as pointing to the fact that just as genes come in slightly different versions, with slightly different effects (holding other factors constant), the same is true of nongenetic factors. Epigenetic markers on genes, due to nutritional environments, litter position and birth order, may also come in slightly different variants, with slightly different effects. Other versions of the claim are much more controversial.

One response to the parity thesis has been to accept the view that genes are just one of a set of individually necessary and collectively sufficient developmental factors, but to argue nonetheless that genes play both a distinctive and especially important role in development (Austin 2015; Lean 2014; Planer 2014). As noted above, perhaps the most promising suggestion along these lines is that genes exert a form of causal control over development that is universal, pervasive and fine-grained. Many features of the phenotypes of every organism exist in an array of somewhat different versions, as a result of allelic variation in causally relevant genes. No other developmental factor exerts control that is similarly universal, pervasive and fine-grained (Woodward 2010; Stegmann 2014). Thus Stegmann illustrates Woodward’s idea that genes exert specific control over phenotypes by contrasting the effects of intervening on, say, the quantity of polymerase on cell activity with intervening on the DNA template itself. Polymerase is critically causally important, but varying its concentration will modify the rate of synthesis, but not the sequences produced. That is not true of modifications of the DNA sequence itself, so the DNA sequence is more causally specific than polymerase.

Shea takes a different approach, arguing that different causal factors have different evolutionary histories. Some causal factors are simply persisting features of the environment (gravity being one example). Others are experienced by the developing organism as a result of histories of selection. Burrows, for example, ensure that eggs and nestlings develop in fairly constant temperature and humidity. But burrows are not naturally selected inheritance mechanisms. They have not come into existence to ensure that a seabird chick resembles its parents. In contrast, some other developmental features are present and act in development because of histories of selection in which the selective advantage is that these mechanisms help ensure parent-offspring similarity. Shea argues that genes, probably epigenetic markings on genes, and perhaps a few other developmental resources are shaped by this form of natural selection. So genes, and perhaps a few other developmental factors, play a distinctive developmental role, even though many other factors are causally necessary (Shea 2011).

In sum then, there are good reasons to be cautious about the use of informational terminology in thinking about development. But it is also possible to over-estimate the strength of the connection between informational conceptions of development and the idea that genes play a uniquely important role in development. There are ways of defending the idea that genes play a special role while acknowledging the interactive character of development. Moreover, an ambitious use of informational concepts is not confined to those within mainstream biological thinking. Eva Jablonka and Marion Lamb defend quite heterodox views of inheritance and evolution, while basing key parts of their work—including an advocacy of “Lamarckian” ideas—around informational concepts (Jablonka and Lamb 2005). They suggest that one of the useful features of informational descriptions is that they allow us to generalize across different heredity systems, comparing their properties in a common currency. In addition, one of the present authors has used informational concepts to distinguish between the evolutionary role of genes from that of other inherited factors whilst demonstrating the evolutionary importance of non-genetic inheritance (Sterelny 2004, 2011). So in various ways, an informational point of view may facilitate discussion of unorthodox theoretical options, including non-genetic mechanisms of inheritance.

Talk of genetic “programs” is common both in popular presentations of biology, and in biology itself. Often, the idea is just a vivid (but perhaps misleading) way of drawing attention to the orderly, well-controlled and highly structured character of development. In its overall results, development is astonishingly stable and predictable, despite the extraordinary complexity of intracellular and intercellular interactions, and despite the fact that the physical context in which development takes place can never be precisely controlled. So when biologists speak, for example, of “programmed cell death”, they could just as well say that in an important class of cases, cell death is predictable, organised, and adaptive.

There are attempts to draw closer and more instructive parallels between computational systems and biological development. In particular, Roger Sansom has made a sustained and detailed attempt to develop close and instructive parallels between biological development and connectionist computational models (Sansom 2008b,a, 2011). This view has the merit of recognising that there is no central control of development; organisms develop as a result of local interactions within and between cells. However, the most promising ideas about program-development parallels seem to us to be ones that point to an apparently close analogy between processes within cells, and the low-level operation of modern computers. One crucial kind of causal process within cells is cascades of up and down-regulation in genetic networks. One gene will make a product that binds to and hence down-regulates another gene, which is then prevented from making a product that up-regulates another… and so on. What we have here is a cascade of events that can often be described in terms of Boolean relationships between variables. One event might only follow from the conjunction of another two, or from a disjunction of them. Down-regulation is a kind of negation, and there can be double and triple negations in a network. Gene regulation networks often have a rich enough structure of this kind for it to be useful to think of them as engaged in a kind of computation. Computer chip “and-gates”, neural “and-gates” and genetic “and-gates” have genuine similarities.

While talking of signalling networks rather than programs, Brett Calcott has shown that positional information in the developing fruitfly embryo depends on this kind of Boolean structure, with limb bud development depending on the cells that differentiate into limb buds integrating one positive with two negative signals, so the buds develop in a regular pattern on the anterior midline of the embryo. Calcott shows that thinking of development in terms of these signalling networks with their Boolean structures has real explanatory value, for it enables us to explain how positional information, for example, can easily be reused in evolution. Wing spots on fruitflies can evolve very easily, for the networks that tell cells where they are on the wing already exist, so the evolution of the wingspot just need a simple mutational change that links that positional information to pigment production (Calcott 2014). Ron Planer agrees that gene regulation has this Boolean structure, and that we can, in effect, represent each gene as instantiating a conditional instruction. The “if” part of the conditional specifies the molecular conditions which turn the gene on; the “then” part of the conditional specifies the amino acid sequence made from the gene. As with Calcott, Planer goes on to point out that these conditional instructions can be and often are, linked together to build complex networks of control. Even so, Planer argues that while these are signalling networks, they should not be thought of as computer programs. For example, the combinations of instructions have no intrinsic order; we can represent each of the genes as a specific conditional instruction, but there is nothing in the set of instructions itself that tells us where to begin and end (Planer 2014).

Information has also become a focus of general discussion of evolutionary processes, especially as they relate to the mechanisms of inheritance. One strand of this discussion misconceives information and its role in biological processes. In particular, G.C. Williams argues that, via reflection on the role of genes in evolution, we can infer that there is an informational “domain” that exists alongside the physical domain of matter and energy (Williams 1992). Richard Dawkins defends a similar view, arguing that the long-term path of evolution is made up of gradual changes in inherited information—as a river that “flows through time, not space” (Dawkins 1995: 4). This is an extension of a more common idea, that there exists such things as “informational genes” that should be understood as distinct from the “material genes” that are made of DNA and localized in space and time (Haig 1997). It is a mistake to think that there are two different things; that there is both a physical entity—a string of bases—and an informational entity, a message. It is true that for evolutionary (and many other) purposes genes are often best thought of in terms of their base sequence (the sequence of C, A, T and G), not in terms of their full set of material properties. This way of thinking is essentially a piece of abstraction (Griesemer 2005). We rightly ignore some properties of DNA and focus on others. But it is a mistake to treat this abstraction as an extra entity, with mysterious relations to the physical domain.

Other ways of linking informational ideas to general issues in evolutionary theory seem more promising. As John Maynard Smith, Eors Szathmáry, Mark Ridley and Richard Dawkins have emphasized in different ways, inheritance mechanisms that give rise to significant evolutionary outcomes must satisfy some rather special conditions (Dawkins 1995; Jablonka and Szathmáry 1995; Szathmáry and Maynard Smith 1997; Ridley 2000). Maynard Smith and Szathmáry claim, for example, that the inheritance system must be unlimited or “indefinite” in its capacity to produce new combinations, but must also maintain high fidelity of transmission. This fact about the relationship between inheritance systems and biological structure is often thought to reveal one of the most pressing problems in explaining the origins of life. If reproduction depends on the replication of a crucial set of ingredients to kick-start the new generation (whether or not we think of those ingredients as instructions), these ingredients must be replicated accurately. Yet accurate replication apparently depends on complex molecular and intracellular machinery that itself is the result of a long regime of adaptive evolution, and hence on deep lineages of living systems. So how could reproduction have begun? (see Ridley 2000 for a thoughtful discussion).

So life itself, the argument goes, depends on the evolution of mechanisms that support a high fidelity flow of information from one generation to the next. More ambitiously still, Maynard Smith and Szathmáry argue that many of the crucial steps in the last four billion years of evolution—their “major transitions in evolution”—involve the creation of new ways of transmitting information across generations—more reliable, more fine-grained, and more powerful ways of making possible the reliable re-creation of form across events of biological reproduction. The transition to a DNA-based inheritance system (probably from a system based on RNA) is one central example. But Maynard Smith and Szathmáry suggest that the transition from great ape forms of social life to human social life is a major transition, in part because of the novel forms of large scale cooperation that typify human social life, but mostly because they see human language as a breakthrough informational technology, revolutionising the possibilities of high fidelity intergenerational cultural learning (MacArthur 1958; Maynard Smith and Szathmáry 1995, 1999).

Maynard Smith and Szathmáry’s work on major transitions has been a landmark in macroevolutionary thinking—in thinking about large scale patterns in the history of life. But the informational dimension of their work has not been taken up. A minor exception is Sterelny (2009). This paper argued that multicelled animal life depended not just on the transmission of more information with high fidelity, but on the control of that information in development, suggesting that the evolution of the egg—a controlled, structured, information-rich developmental environment—was critical to complex animal life. But most focus has been on the other strand of their work on major transitions: on the solution of cooperation problems that then allow previously independent agents to combine into new, more complex agents. Thus in Calcott and Sterelny (2011), none of the papers focused primarily on the expansion or control of intergenerational information flow. That may change. The evolutionarily crucial features of inheritance mechanisms are often now discussed in informational terms, and the combinatorial structure seen in both language and DNA provides a powerful basis for analogical reasoning.

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functionalism | genetics: ecological | genetics: genotype/phenotype distinction | heritability | mental content: teleological theories of | mental representation | naturalism | natural selection | teleology: teleological notions in biology

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bioinformatics

Definition of bioinformatics

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These examples are programmatically compiled from various online sources to illustrate current usage of the word 'bioinformatics.' Any opinions expressed in the examples do not represent those of Merriam-Webster or its editors. Send us feedback about these examples.

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1975, in the meaning defined above

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“Bioinformatics.” Merriam-Webster.com Dictionary , Merriam-Webster, https://www.merriam-webster.com/dictionary/bioinformatics. Accessed 2 May. 2024.

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