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The history of biology traces the study of the living world from
ancient Ancient history is a time period from the beginning of writing and recorded human history to as far as late antiquity. The span of recorded history is roughly 5,000 years, beginning with the Sumerian cuneiform script. Ancient history cove ...
to
modern Modern may refer to: History *Modern history ** Early Modern period ** Late Modern period *** 18th century *** 19th century *** 20th century ** Contemporary history * Moderns, a faction of Freemasonry that existed in the 18th century Philosophy ...
times. Although the concept of ''
biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
'' as a single coherent field arose in the 19th century, the biological sciences emerged from traditions of medicine and natural history reaching back to
Ayurveda Ayurveda () is an alternative medicine system with historical roots in the Indian subcontinent. The theory and practice of Ayurveda is pseudoscientific. Ayurveda is heavily practiced in India and Nepal, where around 80% of the population repor ...
,
ancient Egyptian medicine The medicine of the ancient Egyptians is some of the oldest documented. From the beginnings of the civilization in the late fourth millennium BC until the Persian invasion of 525 BC, Egyptian medical practice went largely unchanged and include ...
and the works of Aristotle and Galen in the ancient Greco-Roman world. This ancient work was further developed in the Middle Ages by Muslim physicians and scholars such as
Avicenna Ibn Sina ( fa, ابن سینا; 980 – June 1037 CE), commonly known in the West as Avicenna (), was a Persian polymath who is regarded as one of the most significant physicians, astronomers, philosophers, and writers of the Islamic ...
. During the European Renaissance and early modern period, biological thought was revolutionized in Europe by a renewed interest in empiricism and the discovery of many novel organisms. Prominent in this movement were Vesalius and
Harvey Harvey, Harveys or Harvey's may refer to: Arts, entertainment, and media * ''Harvey'' (play), a 1944 play by Mary Chase about a man befriended by an invisible anthropomorphic rabbit * Harvey Awards ("Harveys"), one of the most important awards ...
, who used experimentation and careful observation in physiology, and naturalists such as Linnaeus and Buffon who began to classify the diversity of life and the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
, as well as the development and behavior of organisms.
Antonie van Leeuwenhoek Antonie Philips van Leeuwenhoek ( ; ; 24 October 1632 – 26 August 1723) was a Dutch microbiologist and microscopist in the Golden Age of Dutch science and technology. A largely self-taught man in science, he is commonly known as " the ...
revealed by means of
microscopy Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of mic ...
the previously unknown world of microorganisms, laying the groundwork for
cell theory In biology, cell theory is a scientific theory first formulated in the mid-nineteenth century, that living organisms are made up of cells, that they are the basic structural/organizational unit of all organisms, and that all cells come from pre ...
. The growing importance of natural theology, partly a response to the rise of
mechanical philosophy The mechanical philosophy is a form of natural philosophy which compares the universe to a large-scale mechanism (i.e. a machine). The mechanical philosophy is associated with the scientific revolution of early modern Europe. One of the first expo ...
, encouraged the growth of natural history (although it entrenched the
argument from design The teleological argument (from ; also known as physico-theological argument, argument from design, or intelligent design argument) is an argument for the existence of God or, more generally, that complex functionality in the natural world wh ...
). Over the 18th and 19th centuries, biological sciences such as
botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
and zoology became increasingly professional scientific disciplines.
Lavoisier Antoine-Laurent de Lavoisier ( , ; ; 26 August 17438 May 1794),
CNRS (
Alexander von Humboldt Friedrich Wilhelm Heinrich Alexander von Humboldt (14 September 17696 May 1859) was a German polymath, geographer, naturalist, explorer, and proponent of Romantic philosophy and science. He was the younger brother of the Prussian minister, ...
investigated the interaction between organisms and their environment, and the ways this relationship depends on geography—laying the foundations for
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, i ...
, ecology and
ethology Ethology is the scientific study of animal behaviour, usually with a focus on behaviour under natural conditions, and viewing behaviour as an evolutionarily adaptive trait. Behaviourism as a term also describes the scientific and object ...
. Naturalists began to reject essentialism and consider the importance of extinction and the mutability of species.
Cell theory In biology, cell theory is a scientific theory first formulated in the mid-nineteenth century, that living organisms are made up of cells, that they are the basic structural/organizational unit of all organisms, and that all cells come from pre ...
provided a new perspective on the fundamental basis of life. These developments, as well as the results from
embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos an ...
and paleontology, were synthesized in
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
's theory of evolution by natural selection. The end of the 19th century saw the fall of
spontaneous generation Spontaneous generation is a superseded scientific theory that held that living creatures could arise from nonliving matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could arise f ...
and the rise of the
germ theory of disease The germ theory of disease is the currently accepted scientific theory for many diseases. It states that microorganisms known as pathogens or "germs" can lead to disease. These small organisms, too small to be seen without magnification, invade ...
, though the mechanism of
inheritance Inheritance is the practice of receiving private property, titles, debts, entitlements, privileges, rights, and obligations upon the death of an individual. The rules of inheritance differ among societies and have changed over time. Officiall ...
remained a mystery. In the early 20th century, the rediscovery of Mendel's work led to the rapid development of genetics by Thomas Hunt Morgan and his students, and by the 1930s the combination of
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and pop ...
and natural selection in the " neo-Darwinian synthesis". New disciplines developed rapidly, especially after
Watson Watson may refer to: Companies * Actavis, a pharmaceutical company formerly known as Watson Pharmaceuticals * A.S. Watson Group, retail division of Hutchison Whampoa * Thomas J. Watson Research Center, IBM research center * Watson Systems, make ...
and Crick proposed the structure of DNA. Following the establishment of the Central Dogma and the cracking of the
genetic code The genetic code is the set of rules used by living cells to translate information encoded within genetic material ( DNA or RNA sequences of nucleotide triplets, or codons) into proteins. Translation is accomplished by the ribosome, which links ...
, biology was largely split between ''organismal biology''—the fields that deal with whole organisms and groups of organisms—and the fields related to '' cellular and molecular biology''. By the late 20th century, new fields like
genomics Genomics is an interdisciplinary field of biology focusing on the structure, function, evolution, mapping, and editing of genomes. A genome is an organism's complete set of DNA, including all of its genes as well as its hierarchical, three-dim ...
and
proteomics Proteomics is the large-scale study of proteins. Proteins are vital parts of living organisms, with many functions such as the formation of structural fibers of muscle tissue, enzymatic digestion of food, or synthesis and replication of DNA. I ...
were reversing this trend, with organismal biologists using molecular techniques, and molecular and cell biologists investigating the interplay between genes and the environment, as well as the genetics of natural populations of organisms.


Prehistoric times

The earliest humans must have had and passed on knowledge about plants and
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
s to increase their chances of survival. This may have included knowledge of human and animal anatomy and aspects of animal behavior (such as migration patterns). However, the first major turning point in biological knowledge came with the Neolithic Revolution about 10,000 years ago. Humans first domesticated plants for farming, then livestock animals to accompany the resulting sedentary societies.


Earliest roots

In around 3000 to 1200
BCE Common Era (CE) and Before the Common Era (BCE) are year notations for the Gregorian calendar (and its predecessor, the Julian calendar), the world's most widely used calendar era. Common Era and Before the Common Era are alternatives to the o ...
, the Ancient Egyptians and Mesopotamians made contributions to
astronomy Astronomy () is a natural science that studies celestial objects and phenomena. It uses mathematics, physics, and chemistry in order to explain their origin and evolution. Objects of interest include planets, moons, stars, nebulae, galax ...
, mathematics, and medicine, which later entered and shaped Greek natural philosophy of classical antiquity, a period that profoundly influenced the development of what came to be known as biology.


Ancient Egypt

Over a dozen medical papyri have been preserved, most notably the
Edwin Smith Papyrus The Edwin Smith Papyrus is an ancient Egyptian medical text, named after Edwin Smith who bought it in 1862, and the oldest known surgical treatise on trauma. From a cited quotation in another text, it may have been known to ancient surgeons as ...
(the oldest extant surgical handbook) and the Ebers Papyrus (a handbook of preparing and using materia medica for various diseases), both from around 1600 BCE. Ancient Egypt is also known for developing
embalming Embalming is the art and science of preserving human remains by treating them (in its modern form with chemicals) to forestall decomposition. This is usually done to make the deceased suitable for public or private viewing as part of the funeral ...
, which was used for mummification, in order to preserve human remains and forestall decomposition.


Mesopotamia

The Mesopotamians seem to have had little interest in the natural world as such, preferring to study how the gods had ordered the universe.
Animal physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical a ...
was studied for divination, including especially the anatomy of the liver, seen as an important organ in haruspicy. Animal behavior too was studied for divinatory purposes. Most information about the training and domestication of animals was probably transmitted orally, but one text dealing with the training of horses has survived. The ancient Mesopotamians had no distinction between "rational science" and
magic Magic or Magick most commonly refers to: * Magic (supernatural), beliefs and actions employed to influence supernatural beings and forces * Ceremonial magic, encompasses a wide variety of rituals of magic * Magical thinking, the belief that unrel ...
. When a person became ill, doctors prescribed both magical formulas to be recited and medicinal treatments. The earliest medical prescriptions appear in Sumerian during the Third Dynasty of Ur ( 2112 – 2004 BCE). The most extensive Babylonian medical text, however, is the ''Diagnostic Handbook'' written by the ''ummânū'', or chief scholar, Esagil-kin-apli of
Borsippa Borsippa ( Sumerian: BAD.SI.(A).AB.BAKI; Akkadian: ''Barsip'' and ''Til-Barsip'')The Cambridge Ancient History: Prolegomena & Prehistory': Vol. 1, Part 1. Accessed 15 Dec 2010. or Birs Nimrud (having been identified with Nimrod) is an archeolo ...
, during the reign of the Babylonian king
Adad-apla-iddina Adad-apla-iddina, typically inscribed in cuneiform mdIM- DUMU.UŠ-SUM''-na'', mdIM-A-SUM''-na'' or dIM''-ap-lam-i-din-'' 'nam''meaning the storm god “Adad has given me an heir”, was the 8th king of the 2nd Dynasty of Isin and the 4th Dynasty ...
(1069 – 1046 BCE). In East Semitic cultures, the main medicinal authority was an exorcist-healer known as an '' āšipu''. The profession was passed down from father to son and was held in high regard. Of less frequent recourse was the ''asu'', a healer who treated physical symptoms using remedies composed of herbs, animal products, and minerals, as well as potions, enemas, and ointments or
poultices A poultice, also called a cataplasm, is a soft moist mass, often heated and medicated, that is spread on cloth and placed over the skin to treat an aching, inflamed, or painful part of the body. It can be used on wounds, such as cuts. 'Poultice ...
. These physicians, who could be either male or female, also dressed wounds, set limbs, and performed simple surgeries. The ancient Mesopotamians also practiced
prophylaxis Preventive healthcare, or prophylaxis, consists of measures taken for the purposes of disease prevention.Hugh R. Leavell and E. Gurney Clark as "the science and art of preventing disease, prolonging life, and promoting physical and mental hea ...
and took measures to prevent the spread of disease.


Separate developments in China and India

Observations and theories regarding nature and human health, separate from
Western tradition Eugen Joseph Weber (April 24, 1925 – May 17, 2007) was a Romanian-born American historian with a special focus on Western civilization. Weber became a historian because of his interest in politics, an interest dating back to at least the age ...
s, had emerged independently in other civilizations such as those in China and the Indian subcontinent. In ancient China, earlier conceptions can be found dispersed across several different disciplines, including the work of herbologists, physicians, alchemists, and philosophers. The Taoist tradition of
Chinese alchemy Chinese alchemy is an ancient Chinese scientific and technological approach to alchemy, a part of the larger tradition of Taoist / Daoist body-spirit cultivation developed from the traditional Chinese understanding of medicine and the body. Accor ...
, for example, emphasized health (with the ultimate goal being the elixir of life). The system of
classical Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of action ...
usually revolved around the theory of yin and yang, and the
five phases (; Japanese: (); Korean: (); Vietnamese: ''ngũ hành'' (五行)), usually translated as Five Phases or Five Agents, is a fivefold conceptual scheme that many traditional Chinese fields used to explain a wide array of phenomena, from cosmi ...
. Taoist philosophers, such as Zhuangzi in the 4th century BCE, also expressed ideas related to evolution, such as denying the fixity of biological species and speculating that species had developed differing attributes in response to differing environments. One of the oldest organised systems of medicine is known from ancient India in the form of
Ayurveda Ayurveda () is an alternative medicine system with historical roots in the Indian subcontinent. The theory and practice of Ayurveda is pseudoscientific. Ayurveda is heavily practiced in India and Nepal, where around 80% of the population repor ...
, which originated around 1500 BCE from
Atharvaveda The Atharva Veda (, ' from ' and ''veda'', meaning "knowledge") is the "knowledge storehouse of ''atharvāṇas'', the procedures for everyday life".Laurie Patton (2004), Veda and Upanishad, in ''The Hindu World'' (Editors: Sushil Mittal and G ...
(one of the four most ancient books of Indian knowledge, wisdom and culture). The ancient Indian
Ayurveda Ayurveda () is an alternative medicine system with historical roots in the Indian subcontinent. The theory and practice of Ayurveda is pseudoscientific. Ayurveda is heavily practiced in India and Nepal, where around 80% of the population repor ...
tradition independently developed the concept of three humours, resembling that of the
four humours Humorism, the humoral theory, or humoralism, was a system of medicine detailing a supposed makeup and workings of the human body, adopted by Ancient Greek and Roman physicians and philosophers. Humorism began to fall out of favor in the 1850s ...
of
ancient Greek medicine Ancient Greek medicine was a compilation of theories and practices that were constantly expanding through new ideologies and trials. Many components were considered in ancient Greek medicine, intertwining the spiritual with the physical. Specif ...
, though the Ayurvedic system included further complications, such as the body being composed of five elements and seven basic tissues. Ayurvedic writers also classified living things into four categories based on the method of birth (from the womb, eggs, heat & moisture, and seeds) and explained the conception of a fetus in detail. They also made considerable advances in the field of surgery, often without the use of human
dissection Dissection (from Latin ' "to cut to pieces"; also called anatomization) is the dismembering of the body of a deceased animal or plant to study its anatomical structure. Autopsy is used in pathology and forensic medicine to determine the cause o ...
or animal vivisection. One of the earliest Ayurvedic treatises was the '' Sushruta Samhita'', attributed to Sushruta in the 6th century BCE. It was also an early materia medica, describing 700 medicinal plants, 64 preparations from mineral sources, and 57 preparations based on animal sources.


Classical antiquity

The pre-Socratic philosophers asked many questions about life but produced little systematic knowledge of specifically biological interest—though the attempts of the
atomists Atomism (from Greek , ''atomon'', i.e. "uncuttable, indivisible") is a natural philosophy proposing that the physical universe is composed of fundamental indivisible components known as atoms. References to the concept of atomism and its atoms a ...
to explain life in purely physical terms would recur periodically through the history of biology. However, the medical theories of Hippocrates and his followers, especially
humorism Humorism, the humoral theory, or humoralism, was a system of medicine detailing a supposed makeup and workings of the human body, adopted by Ancient Greek and Roman physicians and philosophers. Humorism began to fall out of favor in the 1850s ...
, had a lasting impact. The philosopher Aristotle was the most influential scholar of the living world from classical antiquity. Though his early work in natural philosophy was speculative, Aristotle's later biological writings were more empirical, focusing on biological causation and the diversity of life. He made countless observations of nature, especially the habits and
attributes Attribute may refer to: * Attribute (philosophy), an extrinsic property of an object * Attribute (research), a characteristic of an object * Grammatical modifier, in natural languages * Attribute (computing) In computing, an attribute is a sp ...
of plants and
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
s in the world around him, which he devoted considerable attention to categorizing. In all, Aristotle classified 540 animal species, and dissected at least 50. He believed that intellectual purposes, formal causes, guided all natural processes. Aristotle, and nearly all Western scholars after him until the 18th century, believed that creatures were arranged in a graded scale of perfection rising from plants on up to humans: the ''scala naturae'' or
Great Chain of Being The great chain of being is a hierarchical structure of all matter and life, thought by medieval Christianity to have been decreed by God. The chain begins with God and descends through angels, humans, animals and plants to minerals. The great ...
. Aristotle's successor at the
Lyceum The lyceum is a category of educational institution defined within the education system of many countries, mainly in Europe. The definition varies among countries; usually it is a type of secondary school. Generally in that type of school the ...
, Theophrastus, wrote a series of books on botany—the '' History of Plants''—which survived as the most important contribution of antiquity to botany, even into the Middle Ages. Many of Theophrastus' names survive into modern times, such as ''carpos'' for fruit, and ''pericarpion'' for seed vessel. Dioscorides wrote a pioneering and encyclopaedic pharmacopoeia, ''
De Materia Medica (Latin name for the Greek work , , both meaning "On Medical Material") is a pharmacopoeia of medicinal plants and the medicines that can be obtained from them. The five-volume work was written between 50 and 70 CE by Pedanius Dioscorides, a ...
'', incorporating descriptions of some 600 plants and their uses in medicine. Pliny the Elder, in his '' Natural History'', assembled a similarly encyclopaedic account of things in nature, including accounts of many plants and animals. A few scholars in the
Hellenistic period In Classical antiquity, the Hellenistic period covers the time in Mediterranean history after Classical Greece, between the death of Alexander the Great in 323 BC and the emergence of the Roman Empire, as signified by the Battle of Actium in 3 ...
under the
Ptolemies The Ptolemaic dynasty (; grc, Πτολεμαῖοι, ''Ptolemaioi''), sometimes referred to as the Lagid dynasty (Λαγίδαι, ''Lagidae;'' after Ptolemy I's father, Lagus), was a Macedonian Greek royal dynasty which ruled the Ptolemaic K ...
—particularly
Herophilus of Chalcedon Herophilos (; grc-gre, Ἡρόφιλος; 335–280 BC), sometimes Latinised Herophilus, was a Greek physician regarded as one of the earliest anatomists. Born in Chalcedon, he spent the majority of his life in Alexandria. He was the first sci ...
and Erasistratus of Chios—amended Aristotle's physiological work, even performing dissections and vivisections.
Claudius Galen Aelius Galenus or Claudius Galenus ( el, Κλαύδιος Γαληνός; September 129 – c. AD 216), often Anglicized as Galen () or Galen of Pergamon, was a Greek physician, surgeon and philosopher in the Roman Empire. Considered to be one of ...
became the most important authority on medicine and anatomy. Though a few ancient
atomists Atomism (from Greek , ''atomon'', i.e. "uncuttable, indivisible") is a natural philosophy proposing that the physical universe is composed of fundamental indivisible components known as atoms. References to the concept of atomism and its atoms a ...
such as
Lucretius Titus Lucretius Carus ( , ;  – ) was a Roman poet and philosopher. His only known work is the philosophical poem ''De rerum natura'', a didactic work about the tenets and philosophy of Epicureanism, and which usually is translated into E ...
challenged the teleological Aristotelian viewpoint that all aspects of life are the result of design or purpose, teleology (and after the rise of Christianity, natural theology) would remain central to biological thought essentially until the 18th and 19th centuries.
Ernst W. Mayr Ernst Walter Mayr (; 5 July 1904 – 3 February 2005) was one of the 20th century's leading evolutionary biologists. He was also a renowned taxonomist, tropical explorer, ornithologist, philosopher of biology, and historian of science. His w ...
argued that "Nothing of any real consequence happened in biology after Lucretius and Galen until the Renaissance." The ideas of the Greek traditions of natural history and medicine survived, but they were generally taken unquestioningly in medieval Europe.


Middle Ages

The decline of the
Roman Empire The Roman Empire ( la, Imperium Romanum ; grc-gre, Βασιλεία τῶν Ῥωμαίων, Basileía tôn Rhōmaíōn) was the post-Roman Republic, Republican period of ancient Rome. As a polity, it included large territorial holdings aro ...
led to the disappearance or destruction of much knowledge, though physicians still incorporated many aspects of the Greek tradition into training and practice. In
Byzantium Byzantium () or Byzantion ( grc, Βυζάντιον) was an ancient Greek city in classical antiquity that became known as Constantinople in late antiquity and Istanbul today. The Greek name ''Byzantion'' and its Latinization ''Byzantium'' con ...
and the Islamic world, many of the Greek works were translated into
Arabic Arabic (, ' ; , ' or ) is a Semitic language spoken primarily across the Arab world.Semitic languages: an international handbook / edited by Stefan Weninger; in collaboration with Geoffrey Khan, Michael P. Streck, Janet C. E.Watson; Walte ...
and many of the works of Aristotle were preserved.Mayr, ''The Growth of Biological Thought'', pp 91–94 During the
High Middle Ages The High Middle Ages, or High Medieval Period, was the period of European history that lasted from AD 1000 to 1300. The High Middle Ages were preceded by the Early Middle Ages and were followed by the Late Middle Ages, which ended around AD 1500 ...
, a few European scholars such as
Hildegard of Bingen Hildegard of Bingen (german: Hildegard von Bingen; la, Hildegardis Bingensis; 17 September 1179), also known as Saint Hildegard and the Sibyl of the Rhine, was a German Benedictine abbess and polymath active as a writer, composer, philosopher ...
,
Albertus Magnus Albertus Magnus (c. 1200 – 15 November 1280), also known as Saint Albert the Great or Albert of Cologne, was a German Dominican friar, philosopher, scientist, and bishop. Later canonised as a Catholic saint, he was known during his life ...
and Frederick II wrote on natural history. The rise of European universities, though important for the development of physics and philosophy, had little impact on biological scholarship.


Renaissance

The European Renaissance brought expanded interest in both empirical natural history and physiology. In 1543,
Andreas Vesalius Andreas Vesalius (Latinized from Andries van Wezel) () was a 16th-century anatomist, physician, and author of one of the most influential books on human anatomy, '' De Humani Corporis Fabrica Libri Septem'' (''On the fabric of the human body'' ...
inaugurated the modern era of Western medicine with his seminal
human anatomy The human body is the structure of a human being. It is composed of many different types of cells that together create tissues and subsequently organ systems. They ensure homeostasis and the viability of the human body. It comprises a head, ...
treatise '' De humani corporis fabrica'', which was based on dissection of corpses. Vesalius was the first in a series of anatomists who gradually replaced scholasticism with empiricism in physiology and medicine, relying on first-hand experience rather than authority and abstract reasoning. Via herbalism, medicine was also indirectly the source of renewed empiricism in the study of plants. Otto Brunfels,
Hieronymus Bock Hieronymus Bock ( Latinised Hieronymus Tragus; c. 1498 – 21 February 1554) was a German botanist, physician, and Lutheran minister who began the transition from medieval botany to the modern scientific worldview by arranging plants by their re ...
and Leonhart Fuchs wrote extensively on wild plants, the beginning of a nature-based approach to the full range of plant life.
Bestiaries A bestiary (from ''bestiarum vocabulum'') is a compendium of beasts. Originating in the ancient world, bestiaries were made popular in the Middle Ages in illustrated volumes that described various animals and even rocks. The natural history ...
—a genre that combines both the natural and figurative knowledge of animals—also became more sophisticated, especially with the work of William Turner,
Pierre Belon Pierre Belon (1517–1564) was a French traveller, naturalist, writer and diplomat. Like many others of the Renaissance period, he studied and wrote on a range of topics including ichthyology, ornithology, botany, comparative anatomy, architectu ...
,
Guillaume Rondelet Guillaume Rondelet (27 September 150730 July 1566), also known as Rondeletus/Rondeletius, was Regius professor of medicine at the University of Montpellier in southern France and Chancellor of the University between 1556 and his death in 1566. ...
, Conrad Gessner, and
Ulisse Aldrovandi Ulisse Aldrovandi (11 September 1522 – 4 May 1605) was an Italian naturalist, the moving force behind Bologna's botanical garden, one of the first in Europe. Carl Linnaeus and the comte de Buffon reckoned him the father of natural history ...
. Artists such as
Albrecht Dürer Albrecht Dürer (; ; hu, Ajtósi Adalbert; 21 May 1471 – 6 April 1528),Müller, Peter O. (1993) ''Substantiv-Derivation in Den Schriften Albrecht Dürers'', Walter de Gruyter. . sometimes spelled in English as Durer (without an umlaut) or Due ...
and Leonardo da Vinci, often working with naturalists, were also interested in the bodies of animals and humans, studying physiology in detail and contributing to the growth of anatomical knowledge. The traditions of
alchemy Alchemy (from Arabic: ''al-kīmiyā''; from Ancient Greek: χυμεία, ''khumeía'') is an ancient branch of natural philosophy, a philosophical and protoscientific tradition that was historically practiced in China, India, the Muslim worl ...
and natural magic, especially in the work of
Paracelsus Paracelsus (; ; 1493 – 24 September 1541), born Theophrastus von Hohenheim (full name Philippus Aureolus Theophrastus Bombastus von Hohenheim), was a Swiss physician, alchemist, lay theologian, and philosopher of the German Renaissance. H ...
, also laid claim to knowledge of the living world. Alchemists subjected organic matter to chemical analysis and experimented liberally with both biological and mineral pharmacology. This was part of a larger transition in world views (the rise of the
mechanical philosophy The mechanical philosophy is a form of natural philosophy which compares the universe to a large-scale mechanism (i.e. a machine). The mechanical philosophy is associated with the scientific revolution of early modern Europe. One of the first expo ...
) that continued into the 17th century, as the traditional metaphor of ''nature as organism'' was replaced by the ''nature as machine'' metaphor.


Age of Enlightenment

Systematizing, naming and classifying dominated natural history throughout much of the 17th and 18th centuries. Carl Linnaeus published a basic taxonomy for the natural world in 1735 (variations of which have been in use ever since), and in the 1750s introduced
scientific names In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
for all his species. While Linnaeus conceived of species as unchanging parts of a designed hierarchy, the other great naturalist of the 18th century, Georges-Louis Leclerc, Comte de Buffon, treated species as artificial categories and living forms as malleable—even suggesting the possibility of
common descent Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
. Though he was opposed to evolution, Buffon is a key figure in the
history of evolutionary thought Evolutionary thought, the recognition that species change over time and the perceived understanding of how such processes work, has roots in antiquity—in the ideas of the ancient Greeks, Romans, Chinese, Church Fathers as well as in medieva ...
; his work would influence the evolutionary theories of both Lamarck and Darwin. The discovery and description of new species and the collection of specimens became a passion of scientific gentlemen and a lucrative enterprise for entrepreneurs; many naturalists traveled the globe in search of scientific knowledge and adventure. Extending the work of Vesalius into experiments on still living bodies (of both humans and animals), William Harvey and other natural philosophers investigated the roles of blood, veins and arteries. Harvey's ''
De motu cordis ''Exercitatio Anatomica de Motu Cordis et Sanguinis in Animalibus'' (Latin, 'An Anatomical Exercise on the Motion of the Heart and Blood in Living Beings'), commonly called ''De Motu Cordis'', is the best-known work of the physician William Harv ...
'' in 1628 was the beginning of the end for Galenic theory, and alongside
Santorio Santorio Santorio Santori (29 March, 1561 – 25 February, 1636) also called Santorio Santorio, Santorio de' Sanctoriis, or Sanctorius of Padua and various combinations of these names, was an Italian physiologist, physician, and professor, who introduce ...
's studies of metabolism, it served as an influential model of quantitative approaches to physiology. In the early 17th century, the micro-world of biology was just beginning to open up. A few lensmakers and natural philosophers had been creating crude microscopes since the late 16th century, and Robert Hooke published the seminal '' Micrographia'' based on observations with his own compound microscope in 1665. But it was not until
Antonie van Leeuwenhoek Antonie Philips van Leeuwenhoek ( ; ; 24 October 1632 – 26 August 1723) was a Dutch microbiologist and microscopist in the Golden Age of Dutch science and technology. A largely self-taught man in science, he is commonly known as " the ...
's dramatic improvements in lensmaking beginning in the 1670s—ultimately producing up to 200-fold magnification with a single lens—that scholars discovered
spermatozoa A spermatozoon (; also spelled spermatozoön; ; ) is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote. (A zygote is a single cell, with a complete set of chromosomes, ...
, bacteria,
infusoria Infusoria are minute freshwater life forms including ciliates, euglenoids, protozoa, unicellular algae and small invertebrates. Some authors (e.g., Bütschli) used the term as a synonym for Ciliophora. In modern formal classifications, the term ...
and the sheer strangeness and diversity of microscopic life. Similar investigations by
Jan Swammerdam Jan Swammerdam (February 12, 1637 – February 17, 1680) was a Dutch biologist and microscopist. His work on insects demonstrated that the various phases during the life of an insect— egg, larva, pupa, and adult—are different forms of the ...
led to a new interest in
entomology Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arac ...
and built the basic techniques of microscopic dissection and
staining Staining is a technique used to enhance contrast in samples, generally at the microscopic level. Stains and dyes are frequently used in histology (microscopic study of biological tissues), in cytology (microscopic study of cells), and in the ...
. As the microscopic world was expanding, the macroscopic world was shrinking. Botanists such as John Ray worked to incorporate the flood of newly discovered organisms shipped from across the globe into a coherent taxonomy, and a coherent theology ( natural theology). Debate over another flood, the
Noachian The Noachian is a geologic system and early time period on the planet Mars characterized by high rates of meteorite and asteroid impacts and the possible presence of abundant surface water. The absolute age of the Noachian period is uncertain ...
, catalyzed the development of paleontology; in 1669 Nicholas Steno published an essay on how the remains of living organisms could be trapped in layers of sediment and mineralized to produce fossils. Although Steno's ideas about fossilization were well known and much debated among natural philosophers, an organic origin for all fossils would not be accepted by all naturalists until the end of the 18th century due to philosophical and theological debate about issues such as the age of the earth and extinction.


19th century: the emergence of biological disciplines

Up through the 19th century, the scope of biology was largely divided between medicine, which investigated questions of form and function (i.e., physiology), and natural history, which was concerned with the diversity of life and interactions among different forms of life and between life and non-life. By 1900, much of these domains overlapped, while natural history (and its counterpart natural philosophy) had largely given way to more specialized scientific disciplines—
cytology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living and ...
,
bacteriology Bacteriology is the branch and specialty of biology that studies the morphology, ecology, genetics and biochemistry of bacteria as well as many other aspects related to them. This subdivision of microbiology involves the identification, classificat ...
,
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
,
embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos an ...
, geography, and geology.


Use of the term ''biology''

The term ''biology'' in its modern sense appears to have been introduced independently by
Thomas Beddoes Thomas Beddoes (13 April 176024 December 1808) was an English physician and scientific writer. He was born in Shifnal, Shropshire and died in Bristol fifteen years after opening his medical practice there. He was a reforming practitioner and t ...
(in 1799), Karl Friedrich Burdach (in 1800), Gottfried Reinhold Treviranus (''Biologie oder Philosophie der lebenden Natur'', 1802) and Jean-Baptiste Lamarck (''Hydrogéologie'', 1802). The word itself appears in the title of Volume 3 of Michael Christoph Hanow's ''Philosophiae naturalis sive physicae dogmaticae: Geologia, biologia, phytologia generalis et dendrologia'', published in 1766. Before ''biology,'' there were several terms used for the study of animals and plants. '' Natural history'' referred to the descriptive aspects of biology, though it also included mineralogy and other non-biological fields; from the Middle Ages through the Renaissance, the unifying framework of natural history was the ''scala naturae'' or
Great Chain of Being The great chain of being is a hierarchical structure of all matter and life, thought by medieval Christianity to have been decreed by God. The chain begins with God and descends through angels, humans, animals and plants to minerals. The great ...
. '' Natural philosophy'' and '' natural theology'' encompassed the conceptual and metaphysical basis of plant and animal life, dealing with problems of why organisms exist and behave the way they do, though these subjects also included what is now geology, physics, chemistry, and
astronomy Astronomy () is a natural science that studies celestial objects and phenomena. It uses mathematics, physics, and chemistry in order to explain their origin and evolution. Objects of interest include planets, moons, stars, nebulae, galax ...
. Physiology and (botanical) pharmacology were the province of medicine. ''Botany'', ''Zoology'', and (in the case of fossils) ''Geology'' replaced ''natural history'' and ''natural philosophy'' in the 18th and 19th centuries before ''biology'' was widely adopted. To this day, "botany" and "zoology" are widely used, although they have been joined by other sub-disciplines of biology.


Natural history and natural philosophy

Widespread travel by naturalists in the early-to-mid-19th century resulted in a wealth of new information about the diversity and distribution of living organisms. Of particular importance was the work of
Alexander von Humboldt Friedrich Wilhelm Heinrich Alexander von Humboldt (14 September 17696 May 1859) was a German polymath, geographer, naturalist, explorer, and proponent of Romantic philosophy and science. He was the younger brother of the Prussian minister, ...
, which analyzed the relationship between organisms and their environment (i.e., the domain of natural history) using the quantitative approaches of natural philosophy (i.e., physics and chemistry). Humboldt's work laid the foundations of
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, i ...
and inspired several generations of scientists.


Geology and paleontology

The emerging discipline of geology also brought natural history and natural philosophy closer together; the establishment of the stratigraphic column linked the spatial distribution of organisms to their temporal distribution, a key precursor to concepts of evolution. Georges Cuvier and others made great strides in
comparative anatomy Comparative anatomy is the study of similarities and differences in the anatomy of different species. It is closely related to evolutionary biology and phylogeny (the evolution of species). The science began in the classical era, continuing in ...
and paleontology in the late 1790s and early 19th century. In a series of lectures and papers that made detailed comparisons between living mammals and fossil remains Cuvier was able to establish that the fossils were remains of species that had become extinct—rather than being remains of species still alive elsewhere in the world, as had been widely believed. Fossils discovered and described by Gideon Mantell, William Buckland, Mary Anning, and
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. ...
among others helped establish that there had been an 'age of reptiles' that had preceded even the prehistoric mammals. These discoveries captured the public imagination and focused attention on the history of life on earth. Most of these geologists held to
catastrophism In geology, catastrophism theorises that the Earth has largely been shaped by sudden, short-lived, violent events, possibly worldwide in scope. This contrasts with uniformitarianism (sometimes called gradualism), according to which slow incremen ...
, but
Charles Lyell Sir Charles Lyell, 1st Baronet, (14 November 1797 – 22 February 1875) was a Scottish geologist who demonstrated the power of known natural causes in explaining the earth's history. He is best known as the author of ''Principles of Geolo ...
's influential ''Principles of Geology'' (1830) popularised Hutton's
uniformitarianism Uniformitarianism, also known as the Doctrine of Uniformity or the Uniformitarian Principle, is the assumption that the same natural laws and processes that operate in our present-day scientific observations have always operated in the universe in ...
, a theory that explained the geological past and present on equal terms.


Evolution and biogeography

The most significant evolutionary theory before Darwin's was that of Jean-Baptiste Lamarck; based on the inheritance of acquired characteristics (an inheritance mechanism that was widely accepted until the 20th century), it described a chain of development stretching from the lowliest microbe to humans. The British naturalist
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
, combining the biogeographical approach of Humboldt, the uniformitarian geology of Lyell, Thomas Malthus's writings on population growth, and his own morphological expertise, created a more successful evolutionary theory based on natural selection; similar evidence led
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was a British naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution through natural sele ...
to independently reach the same conclusions. The 1859 publication of Darwin's theory in '' On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life'' is often considered the central event in the history of modern biology. Darwin's established credibility as a naturalist, the sober tone of the work, and most of all the sheer strength and volume of evidence presented, allowed ''Origin'' to succeed where previous evolutionary works such as the anonymous '' Vestiges of Creation'' had failed. Most scientists were convinced of evolution and
common descent Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
by the end of the 19th century. However, natural selection would not be accepted as the primary mechanism of evolution until well into the 20th century, as most contemporary theories of heredity seemed incompatible with the inheritance of random variation. Wallace, following on earlier work by
de Candolle Augustin Pyramus (or Pyrame) de Candolle (, , ; 4 February 17789 September 1841) was a Swiss botanist. René Louiche Desfontaines launched de Candolle's botanical career by recommending him at a herbarium. Within a couple of years de Candolle ...
, Humboldt and Darwin, made major contributions to zoogeography. Because of his interest in the transmutation hypothesis, he paid particular attention to the geographical distribution of closely allied species during his field work first in South America and then in the Malay archipelago. While in the archipelago he identified the Wallace line, which runs through the
Spice Islands A spice is a seed, fruit, root, bark, or other plant substance primarily used for flavoring or coloring food. Spices are distinguished from herbs, which are the leaves, flowers, or stems of plants used for flavoring or as a garnish. Spices ar ...
dividing the fauna of the archipelago between an Asian zone and a New Guinea/Australian zone. His key question, as to why the fauna of islands with such similar climates should be so different, could only be answered by considering their origin. In 1876 he wrote ''The Geographical Distribution of Animals'', which was the standard reference work for over half a century, and a sequel, ''Island Life'', in 1880 that focused on island biogeography. He extended the six-zone system developed by
Philip Sclater Philip Lutley Sclater (4 November 1829 – 27 June 1913) was an English lawyer and zoologist. In zoology, he was an expert ornithologist, and identified the main zoogeographic regions of the world. He was Secretary of the Zoological Society ...
for describing the geographical distribution of birds to animals of all kinds. His method of tabulating data on animal groups in geographic zones highlighted the discontinuities; and his appreciation of evolution allowed him to propose rational explanations, which had not been done before. The scientific study of heredity grew rapidly in the wake of Darwin's ''Origin of Species'' with the work of Francis Galton and the biometricians. The origin of genetics is usually traced to the 1866 work of the monk Gregor Mendel, who would later be credited with the
laws of inheritance Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later popularize ...
. However, his work was not recognized as significant until 35 years afterward. In the meantime, a variety of theories of inheritance (based on
pangenesis Pangenesis was Charles Darwin's hypothetical mechanism for heredity, in which he proposed that each part of the body continually emitted its own type of small organic particles called gemmules that aggregated in the gonads, contributing heritabl ...
, orthogenesis, or other mechanisms) were debated and investigated vigorously.
Embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos an ...
and ecology also became central biological fields, especially as linked to evolution and popularized in the work of Ernst Haeckel. Most of the 19th century work on heredity, however, was not in the realm of natural history, but that of experimental physiology.


Physiology

Over the course of the 19th century, the scope of physiology expanded greatly, from a primarily medically oriented field to a wide-ranging investigation of the physical and chemical processes of life—including plants, animals, and even microorganisms in addition to man. ''Living things as machines'' became a dominant metaphor in biological (and social) thinking.


Cell theory, embryology and germ theory

Advances in
microscopy Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of mic ...
also had a profound impact on biological thinking. In the early 19th century, a number of biologists pointed to the central importance of the
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
. In 1838 and 1839, Schleiden and Schwann began promoting the ideas that (1) the basic unit of organisms is the cell and (2) that individual cells have all the characteristics of life, though they opposed the idea that (3) all cells come from the division of other cells. Thanks to the work of
Robert Remak Robert Remak (26 July 1815 – 29 August 1865) was a Jewish Polish-German embryologist, physiologist, and neurologist, born in Posen, Prussia, who discovered that the origin of cells was by the division of pre-existing cells. as well as several ...
and Rudolf Virchow, however, by the 1860s most biologists accepted all three tenets of what came to be known as
cell theory In biology, cell theory is a scientific theory first formulated in the mid-nineteenth century, that living organisms are made up of cells, that they are the basic structural/organizational unit of all organisms, and that all cells come from pre ...
. Cell theory led biologists to re-envision individual organisms as interdependent assemblages of individual cells. Scientists in the rising field of
cytology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living and ...
, armed with increasingly powerful microscopes and new
staining Staining is a technique used to enhance contrast in samples, generally at the microscopic level. Stains and dyes are frequently used in histology (microscopic study of biological tissues), in cytology (microscopic study of cells), and in the ...
methods, soon found that even single cells were far more complex than the homogeneous fluid-filled chambers described by earlier microscopists. Robert Brown had described the
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: * Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucl ...
in 1831, and by the end of the 19th century cytologists identified many of the key cell components: chromosomes,
centrosome In cell biology, the centrosome (Latin centrum 'center' + Greek sōma 'body') (archaically cytocentre) is an organelle that serves as the main microtubule organizing center (MTOC) of the animal cell, as well as a regulator of cell-cycle progr ...
s mitochondria,
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s, and other structures made visible through staining. Between 1874 and 1884
Walther Flemming Walther Flemming (21 April 1843 – 4 August 1905) was a German biologist and a founder of cytogenetics. He was born in Sachsenberg (now part of Schwerin) as the fifth child and only son of the psychiatrist Carl Friedrich Flemming (1799–1880 ...
described the discrete stages of mitosis, showing that they were not artifacts of staining but occurred in living cells, and moreover, that chromosomes doubled in number just before the cell divided and a daughter cell was produced. Much of the research on cell reproduction came together in
August Weismann August Friedrich Leopold Weismann FRS (For), HonFRSE, LLD (17 January 18345 November 1914) was a German evolutionary biologist. Fellow German Ernst Mayr ranked him as the second most notable evolutionary theorist of the 19th century, after Cha ...
's theory of heredity: he identified the nucleus (in particular chromosomes) as the hereditary material, proposed the distinction between
somatic cell A somatic cell (from Ancient Greek σῶμα ''sôma'', meaning "body"), or vegetal cell, is any biological cell forming the body of a multicellular organism other than a gamete, germ cell, gametocyte or undifferentiated stem cell. Such cells comp ...
s and
germ cell Germ or germs may refer to: Science * Germ (microorganism), an informal word for a pathogen * Germ cell, cell that gives rise to the gametes of an organism that reproduces sexually * Germ layer, a primary layer of cells that forms during embr ...
s (arguing that chromosome number must be halved for germ cells, a precursor to the concept of meiosis), and adopted Hugo de Vries's theory of pangenes. Weismannism was extremely influential, especially in the new field of experimental
embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos an ...
. By the mid-1850s the miasma theory of disease was largely superseded by the
germ theory of disease The germ theory of disease is the currently accepted scientific theory for many diseases. It states that microorganisms known as pathogens or "germs" can lead to disease. These small organisms, too small to be seen without magnification, invade ...
, creating extensive interest in microorganisms and their interactions with other forms of life. By the 1880s,
bacteriology Bacteriology is the branch and specialty of biology that studies the morphology, ecology, genetics and biochemistry of bacteria as well as many other aspects related to them. This subdivision of microbiology involves the identification, classificat ...
was becoming a coherent discipline, especially through the work of
Robert Koch Heinrich Hermann Robert Koch ( , ; 11 December 1843 – 27 May 1910) was a German physician and microbiologist. As the discoverer of the specific causative agents of deadly infectious diseases including tuberculosis, cholera (though the bacteri ...
, who introduced methods for growing pure cultures on agar gels containing specific nutrients in Petri dishes. The long-held idea that living organisms could easily originate from nonliving matter (
spontaneous generation Spontaneous generation is a superseded scientific theory that held that living creatures could arise from nonliving matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could arise f ...
) was attacked in a series of experiments carried out by Louis Pasteur, while debates over vitalism vs. mechanism (a perennial issue since the time of Aristotle and the Greek atomists) continued apace.


Rise of organic chemistry and experimental physiology

In chemistry, one central issue was the distinction between organic and inorganic substances, especially in the context of organic transformations such as fermentation and putrefaction. Since Aristotle these had been considered essentially biological ('' vital'') processes. However,
Friedrich Wöhler Friedrich Wöhler () FRS(For) HonFRSE (31 July 180023 September 1882) was a German chemist known for his work in inorganic chemistry, being the first to isolate the chemical elements beryllium and yttrium in pure metallic form. He was the first ...
,
Justus Liebig Justus Freiherr von Liebig (12 May 1803 – 20 April 1873) was a German scientist who made major contributions to agricultural and biological chemistry, and is considered one of the principal founders of organic chemistry. As a professor at the ...
and other pioneers of the rising field of organic chemistry—building on the work of Lavoisier—showed that the organic world could often be analyzed by physical and chemical methods. In 1828 Wöhler showed that the organic substance
urea Urea, also known as carbamide, is an organic compound with chemical formula . This amide has two amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest amide of carbamic acid. Urea serves an important ...
could be created by chemical means that do not involve life, providing a powerful challenge to vitalism. Cell extracts ("ferments") that could effect chemical transformations were discovered, beginning with
diastase A diastase (; from Greek διάστασις, "separation") is any one of a group of enzymes that catalyses the breakdown of starch into maltose. Alpha amylase degrades starch to a mixture of the disaccharide maltose; the trisaccharide maltotriose, ...
in 1833. By the end of the 19th century the concept of
enzymes Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. ...
was well established, though equations of
chemical kinetics Chemical kinetics, also known as reaction kinetics, is the branch of physical chemistry that is concerned with understanding the rates of chemical reactions. It is to be contrasted with chemical thermodynamics, which deals with the direction in wh ...
would not be applied to enzymatic reactions until the early 20th century. Physiologists such as
Claude Bernard Claude Bernard (; 12 July 1813 – 10 February 1878) was a French physiologist. Historian I. Bernard Cohen of Harvard University called Bernard "one of the greatest of all men of science". He originated the term '' milieu intérieur'', and the ...
explored (through vivisection and other experimental methods) the chemical and physical functions of living bodies to an unprecedented degree, laying the groundwork for endocrinology (a field that developed quickly after the discovery of the first hormone, secretin, in 1902),
biomechanics Biomechanics is the study of the structure, function and motion of the mechanical aspects of biological systems, at any level from whole organisms to organs, cells and cell organelles, using the methods of mechanics. Biomechanics is a branch of ...
, and the study of nutrition and
digestion Digestion is the breakdown of large insoluble food molecules into small water-soluble food molecules so that they can be absorbed into the watery blood plasma. In certain organisms, these smaller substances are absorbed through the small intes ...
. The importance and diversity of experimental physiology methods, within both medicine and biology, grew dramatically over the second half of the 19th century. The control and manipulation of life processes became a central concern, and experiment was placed at the center of biological education.


Twentieth century biological sciences

At the beginning of the 20th century, biological research was largely a professional endeavour. Most work was still done in the natural history mode, which emphasized morphological and phylogenetic analysis over experiment-based causal explanations. However, anti- vitalist experimental physiologists and embryologists, especially in Europe, were increasingly influential. The tremendous success of experimental approaches to development, heredity, and metabolism in the 1900s and 1910s demonstrated the power of experimentation in biology. In the following decades, experimental work replaced natural history as the dominant mode of research.


Ecology and environmental science

In the early 20th century, naturalists were faced with increasing pressure to add rigor and preferably experimentation to their methods, as the newly prominent laboratory-based biological disciplines had done. Ecology had emerged as a combination of biogeography with the
biogeochemical cycle A biogeochemical cycle (or more generally a cycle of matter) is the pathway by which a chemical substance cycles (is turned over or moves through) the biotic and the abiotic compartments of Earth. The biotic compartment is the biosphere and the ...
concept pioneered by chemists; field biologists developed quantitative methods such as the quadrat and adapted laboratory instruments and cameras for the field to further set their work apart from traditional natural history. Zoologists and botanists did what they could to mitigate the unpredictability of the living world, performing laboratory experiments and studying semi-controlled natural environments such as gardens; new institutions like the Carnegie Station for Experimental Evolution and the Marine Biological Laboratory provided more controlled environments for studying organisms through their entire life cycles. The
ecological succession Ecological succession is the process of change in the species structure of an ecological community over time. The time scale can be decades (for example, after a wildfire) or more or less. Bacteria allows for the cycling of nutrients such as ca ...
concept, pioneered in the 1900s and 1910s by Henry Chandler Cowles and
Frederic Clements Frederic Edward Clements (September 16, 1874 – July 26, 1945) was an American plant ecologist and pioneer in the study of plant ecology and vegetation succession. Biography Born in Lincoln, Nebraska, he studied botany at the University of Neb ...
, was important in early plant ecology. Alfred Lotka's predator-prey equations,
G. Evelyn Hutchinson George Evelyn Hutchinson (January 30, 1903 – May 17, 1991) was a British ecologist sometimes described as the "father of modern ecology." He contributed for more than sixty years to the fields of limnology, systems ecology, radiation ecolog ...
's studies of the biogeography and biogeochemical structure of lakes and rivers ( limnology) and Charles Elton's studies of animal
food chain A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), det ...
s were pioneers among the succession of quantitative methods that colonized the developing ecological specialties. Ecology became an independent discipline in the 1940s and 1950s after
Eugene P. Odum Eugene Pleasants Odum (September 17, 1913 – August 10, 2002) was an American biologist at the University of Georgia known for his pioneering work on ecosystem ecology. He and his brother Howard T. Odum wrote the popular ecology textbook, ''Funda ...
synthesized many of the concepts of
ecosystem ecology Ecosystem ecology is the integrated study of living ( biotic) and non-living (abiotic) components of ecosystems and their interactions within an ecosystem framework. This science examines how ecosystems work and relates this to their components ...
, placing relationships between groups of organisms (especially material and energy relationships) at the center of the field. In the 1960s, as evolutionary theorists explored the possibility of multiple units of selection, ecologists turned to evolutionary approaches. In population ecology, debate over group selection was brief but vigorous; by 1970, most biologists agreed that natural selection was rarely effective above the level of individual organisms. The evolution of ecosystems, however, became a lasting research focus. Ecology expanded rapidly with the rise of the environmental movement; the
International Biological Program The International Biological Program (IBP) was an effort between 1964 and 1974 to coordinate large-scale ecological and environmental studies. Organized in the wake of the successful International Geophysical Year (IGY) of 1957-1958, the Internati ...
attempted to apply the methods of big science (which had been so successful in the physical sciences) to ecosystem ecology and pressing environmental issues, while smaller-scale independent efforts such as
island biogeography Insular biogeography or island biogeography is a field within biogeography that examines the factors that affect the species richness and diversification of isolated natural communities. The theory was originally developed to explain the pattern of ...
and the
Hubbard Brook Experimental Forest Hubbard Brook Experimental Forest is an area of land in the towns of Woodstock, Ellsworth and Thornton in the White Mountains of New Hampshire that functions as an outdoor laboratory for ecological studies. It was initially established in 1955 ...
helped redefine the scope of an increasingly diverse discipline.


Classical genetics, the modern synthesis, and evolutionary theory

1900 marked the so-called ''rediscovery of Mendel'': Hugo de Vries,
Carl Correns Carl Erich Correns (19 September 1864 – 14 February 1933) was a German botanist and geneticist notable primarily for his independent discovery of the principles of heredity, which he achieved simultaneously but independently of the botanist ...
, and Erich von Tschermak independently arrived at
Mendel's laws Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later populariz ...
(which were not actually present in Mendel's work). Soon after, cytologists (cell biologists) proposed that chromosomes were the hereditary material. Between 1910 and 1915, Thomas Hunt Morgan and the " Drosophilists" in his fly lab forged these two ideas—both controversial—into the "Mendelian-chromosome theory" of heredity. They quantified the phenomenon of genetic linkage and postulated that genes reside on chromosomes like beads on string; they hypothesized crossing over to explain linkage and constructed
genetic map Genetic linkage is the tendency of DNA sequences that are close together on a chromosome to be inherited together during the meiosis phase of sexual reproduction. Two genetic markers that are physically near to each other are unlikely to be sepa ...
s of the fruit fly '' Drosophila melanogaster'', which became a widely used model organism. Hugo de Vries tried to link the new genetics with evolution; building on his work with heredity and
hybridization Hybridization (or hybridisation) may refer to: *Hybridization (biology), the process of combining different varieties of organisms to create a hybrid *Orbital hybridization, in chemistry, the mixing of atomic orbitals into new hybrid orbitals *Nu ...
, he proposed a theory of
mutationism Mutationism is one of several alternatives to evolution by natural selection that have existed both before and after the publication of Charles Darwin's 1859 book ''On the Origin of Species''. In the theory, mutation was the source of novelty, c ...
, which was widely accepted in the early 20th century. Lamarckism, or the theory of inheritance of acquired characteristics also had many adherents. Darwinism was seen as incompatible with the continuously variable traits studied by biometricians, which seemed only partially heritable. In the 1920s and 1930s—following the acceptance of the Mendelian-chromosome theory— the emergence of the discipline of
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and pop ...
, with the work of
R.A. Fisher Sir Ronald Aylmer Fisher (17 February 1890 – 29 July 1962) was a British polymath who was active as a mathematician, statistician, biologist, geneticist, and academic. For his work in statistics, he has been described as "a genius who a ...
, J.B.S. Haldane and Sewall Wright, unified the idea of evolution by natural selection with
Mendelian genetics Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later popularized ...
, producing the modern synthesis. The
inheritance of acquired characters Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism, is the notion that an organism can pass on to its offspring physical characteristics that the parent organism acquired through use or disuse during its lifetime. It is also calle ...
was rejected, while mutationism gave way as genetic theories matured. In the second half of the century the ideas of population genetics began to be applied in the new discipline of the genetics of behavior, sociobiology, and, especially in humans,
evolutionary psychology Evolutionary psychology is a theoretical approach in psychology that examines cognition and behavior from a modern evolutionary perspective. It seeks to identify human psychological adaptations with regards to the ancestral problems they evolv ...
. In the 1960s W.D. Hamilton and others developed
game theory Game theory is the study of mathematical models of strategic interactions among rational agents. Myerson, Roger B. (1991). ''Game Theory: Analysis of Conflict,'' Harvard University Press, p.&nbs1 Chapter-preview links, ppvii–xi It has applic ...
approaches to explain
altruism Altruism is the principle and moral practice of concern for the welfare and/or happiness of other human beings or animals, resulting in a quality of life both material and spiritual. It is a traditional virtue in many cultures and a core as ...
from an evolutionary perspective through kin selection. The possible origin of higher organisms through endosymbiosis, and contrasting approaches to molecular evolution in the gene-centered view (which held selection as the predominant cause of evolution) and the neutral theory (which made genetic drift a key factor) spawned perennial debates over the proper balance of
adaptationism Adaptationism (also known as functionalism) is the Darwinian view that many physical and psychological traits of organisms are evolved adaptations. Pan-adaptationism is the strong form of this, deriving from the early 20th century modern synthes ...
and contingency in evolutionary theory. In the 1970s Stephen Jay Gould and Niles Eldredge proposed the theory of punctuated equilibrium which holds that stasis is the most prominent feature of the fossil record, and that most evolutionary changes occur rapidly over relatively short periods of time. In 1980 Luis Alvarez and
Walter Alvarez Walter Alvarez (born October 3, 1940) is a professor in the Earth and Planetary Science department at the University of California, Berkeley. He is most widely known for the theory that dinosaurs were killed by an asteroid impact, developed in co ...
proposed the hypothesis that an impact event was responsible for the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With th ...
. Also in the early 1980s, statistical analysis of the fossil record of marine organisms published by
Jack Sepkoski Joseph John Sepkoski Jr. (July 26, 1948 – May 1, 1999) was a University of Chicago paleontologist. Sepkoski studied the fossil record and the diversity of life on Earth. Sepkoski and David Raup contributed to the knowledge of extinction events. ...
and
David M. Raup David M. Raup (April 24, 1933 – July 9, 2015) was a University of Chicago paleontologist. Raup studied the fossil record and the diversity of life on Earth. Raup contributed to the knowledge of extinction events along with his colleague Jack S ...
led to a better appreciation of the importance of mass extinction events to the history of life on earth.


Biochemistry, microbiology, and molecular biology

By the end of the 19th century all of the major pathways of drug metabolism had been discovered, along with the outlines of protein and fatty acid metabolism and urea synthesis. In the early decades of the 20th century, the minor components of foods in human nutrition, the
vitamins A vitamin is an organic molecule (or a set of molecules closely related chemically, i.e. vitamers) that is an essential micronutrient that an organism needs in small quantities for the proper functioning of its metabolism. Essential nutri ...
, began to be isolated and synthesized. Improved laboratory techniques such as chromatography and electrophoresis led to rapid advances in physiological chemistry, which—as ''biochemistry''—began to achieve independence from its medical origins. In the 1920s and 1930s, biochemists—led by Hans Krebs and
Carl Carl may refer to: * Carl, Georgia, city in USA * Carl, West Virginia, an unincorporated community *Carl (name), includes info about the name, variations of the name, and a list of people with the name * Carl², a TV series * "Carl", an episode of ...
and
Gerty Cori Gerty Theresa Cori (; August 15, 1896 – October 26, 1957) was an Austro-Hungarian and American biochemist who in 1947 was the third woman to win a Nobel Prize in science, and the first woman to be awarded the Nobel Prize in Physiology or Me ...
—began to work out many of the central
metabolic pathways In biochemistry, a metabolic pathway is a linked series of chemical reactions occurring within a cell. The reactants, products, and intermediates of an enzymatic reaction are known as metabolites, which are modified by a sequence of chemical reac ...
of life: the
citric acid cycle The citric acid cycle (CAC)—also known as the Krebs cycle or the TCA cycle (tricarboxylic acid cycle)—is a series of chemical reactions to release stored energy through the oxidation of acetyl-CoA derived from carbohydrates, fats, and protei ...
, glycogenesis and
glycolysis Glycolysis is the metabolic pathway that converts glucose () into pyruvate (). The free energy released in this process is used to form the high-energy molecules adenosine triphosphate (ATP) and reduced nicotinamide adenine dinucleotide (NADH) ...
, and the synthesis of
steroid A steroid is a biologically active organic compound with four rings arranged in a specific molecular configuration. Steroids have two principal biological functions: as important components of cell membranes that alter membrane fluidity; and ...
s and porphyrins. Between the 1930s and 1950s,
Fritz Lipmann Fritz Albert Lipmann (; June 12, 1899 – July 24, 1986) was a German-American biochemist and a co-discoverer in 1945 of coenzyme A. For this, together with other research on coenzyme A, he was awarded the Nobel Prize in Physiology or Medicine in ...
and others established the role of ATP as the universal carrier of energy in the cell, and mitochondria as the powerhouse of the cell. Such traditionally biochemical work continued to be very actively pursued throughout the 20th century and into the 21st.


Origins of molecular biology

Following the rise of classical genetics, many biologists—including a new wave of physical scientists in biology—pursued the question of the gene and its physical nature. Warren Weaver—head of the science division of the Rockefeller Foundation—issued grants to promote research that applied the methods of physics and chemistry to basic biological problems, coining the term '' molecular biology'' for this approach in 1938; many of the significant biological breakthroughs of the 1930s and 1940s were funded by the Rockefeller Foundation. Like biochemistry, the overlapping disciplines of
bacteriology Bacteriology is the branch and specialty of biology that studies the morphology, ecology, genetics and biochemistry of bacteria as well as many other aspects related to them. This subdivision of microbiology involves the identification, classificat ...
and virology (later combined as ''microbiology''), situated between science and medicine, developed rapidly in the early 20th century.
Félix d'Herelle Felix may refer to: * Felix (name), people and fictional characters with the name Places * Arabia Felix is the ancient Latin name of Yemen * Felix, Spain, a municipality of the province Almería, in the autonomous community of Andalusia, ...
's isolation of
bacteriophage A bacteriophage (), also known informally as a ''phage'' (), is a duplodnaviria virus that infects and replicates within bacteria and archaea. The term was derived from "bacteria" and the Greek φαγεῖν ('), meaning "to devour". Bacteri ...
during World War I initiated a long line of research focused on phage viruses and the bacteria they infect. The development of standard, genetically uniform organisms that could produce repeatable experimental results was essential for the development of
molecular genetics Molecular genetics is a sub-field of biology that addresses how differences in the structures or expression of DNA molecules manifests as variation among organisms. Molecular genetics often applies an "investigative approach" to determine the ...
. After early work with ''Drosophila'' and maize, the adoption of simpler model systems like the bread mold ''
Neurospora crassa ''Neurospora crassa'' is a type of red bread mold of the phylum Ascomycota. The genus name, meaning "nerve spore" in Greek, refers to the characteristic striations on the spores. The first published account of this fungus was from an infestation ...
'' made it possible to connect genetics to biochemistry, most importantly with
Beadle A beadle, sometimes spelled bedel, is an official of a church or synagogue who may usher, keep order, make reports, and assist in religious functions; or a minor official who carries out various civil, educational, or ceremonial duties on the ...
and Tatum's
one gene-one enzyme hypothesis 1 (one, unit, unity) is a number representing a single or the only entity. 1 is also a numerical digit and represents a single unit of counting or measurement. For example, a line segment of ''unit length'' is a line segment of length 1. ...
in 1941. Genetics experiments on even simpler systems like
tobacco mosaic virus ''Tobacco mosaic virus'' (TMV) is a positive-sense single-stranded RNA virus species in the genus '' Tobamovirus'' that infects a wide range of plants, especially tobacco and other members of the family Solanaceae. The infection causes character ...
and
bacteriophage A bacteriophage (), also known informally as a ''phage'' (), is a duplodnaviria virus that infects and replicates within bacteria and archaea. The term was derived from "bacteria" and the Greek φαγεῖν ('), meaning "to devour". Bacteri ...
, aided by the new technologies of
electron microscopy An electron microscope is a microscope that uses a beam of accelerated electrons as a source of illumination. As the wavelength of an electron can be up to 100,000 times shorter than that of visible light photons, electron microscopes have a hi ...
and ultracentrifugation, forced scientists to re-evaluate the literal meaning of ''life''; virus heredity and reproducing nucleoprotein cell structures outside the nucleus ("plasmagenes") complicated the accepted Mendelian-chromosome theory.
Oswald Avery Oswald Theodore Avery Jr. (October 21, 1877 – February 20, 1955) was a Canadian-American physician and medical researcher. The major part of his career was spent at the Rockefeller Hospital in New York City. Avery was one of the first molecul ...
showed in 1943 that DNA was likely the genetic material of the chromosome, not its protein; the issue was settled decisively with the 1952
Hershey–Chase experiment The Hershey–Chase experiments were a series of experiments conducted in 1952 by Alfred Hershey and Martha Chase that helped to confirm that DNA is genetic material. While DNA had been known to biologists since 1869, many scientists still as ...
—one of many contributions from the so-called phage group centered around physicist-turned-biologist
Max Delbrück Max Ludwig Henning Delbrück (; September 4, 1906 – March 9, 1981) was a German–American biophysicist who participated in launching the molecular biology research program in the late 1930s. He stimulated physical scientists' interest int ...
. In 1953 James Watson and Francis Crick, building on the work of
Maurice Wilkins Maurice Hugh Frederick Wilkins (15 December 1916 – 5 October 2004) was a New Zealand-born British biophysicist and Nobel laureate whose research spanned multiple areas of physics and biophysics, contributing to the scientific understanding ...
and Rosalind Franklin, suggested that the structure of DNA was a double helix. In their famous paper "
Molecular structure of Nucleic Acids "Molecular Structure of Nucleic Acids: A Structure for Deoxyribose Nucleic Acid" was the first article published to describe the discovery of the double helix structure of DNA, using X-ray diffraction and the mathematics of a helix transform. I ...
", Watson and Crick noted coyly, "It has not escaped our notice that the specific pairing we have postulated immediately suggests a possible copying mechanism for the genetic material." After the 1958 Meselson–Stahl experiment confirmed the
semiconservative replication Semiconservative replication describe the mechanism of DNA replication in all known cells. DNA replication occurs on multiple origins of replication along the DNA template strand (antinsense strand). As the DNA double helix is unwound by helicase, ...
of DNA, it was clear to most biologists that nucleic acid sequence must somehow determine
amino acid sequence Protein primary structure is the linear sequence of amino acids in a peptide or protein. By convention, the primary structure of a protein is reported starting from the amino-terminal (N) end to the carboxyl-terminal (C) end. Protein biosynthes ...
in proteins; physicist
George Gamow George Gamow (March 4, 1904 – August 19, 1968), born Georgiy Antonovich Gamov ( uk, Георгій Антонович Гамов, russian: Георгий Антонович Гамов), was a Russian-born Soviet and American polymath, theoret ...
proposed that a fixed
genetic code The genetic code is the set of rules used by living cells to translate information encoded within genetic material ( DNA or RNA sequences of nucleotide triplets, or codons) into proteins. Translation is accomplished by the ribosome, which links ...
connected proteins and DNA. Between 1953 and 1961, there were few known biological sequences—either DNA or protein—but an abundance of proposed code systems, a situation made even more complicated by expanding knowledge of the intermediate role of RNA. In 1961, it was demonstrated that when a gene encodes a protein, three sequential bases of a gene’s DNA specify each successive amino acid of the protein. Thus the
genetic code The genetic code is the set of rules used by living cells to translate information encoded within genetic material ( DNA or RNA sequences of nucleotide triplets, or codons) into proteins. Translation is accomplished by the ribosome, which links ...
is a triplet code, where each triplet (called a codon) specifies a particular amino acid. Furthermore, it was shown that the codons do not overlap with each other in the DNA sequence encoding a protein, and that each sequence is read from a fixed starting point. To actually decipher the code, it took an extensive series of experiments in biochemistry and bacterial genetics, between 1961 and 1966—most importantly the work of Nirenberg and Khorana. During 1962-1964, numerous conditional lethal mutants of a bacterial virus were isolated. These mutants were used in several different labs to advance fundamental understanding of the functions and interactions of the proteins employed in the machinery of DNA replication, DNA repair, DNA recombination, and in the assembly of molecular structures.


Expansion of molecular biology

In addition to the Division of Biology at
Caltech The California Institute of Technology (branded as Caltech or CIT)The university itself only spells its short form as "Caltech"; the institution considers other spellings such a"Cal Tech" and "CalTech" incorrect. The institute is also occasional ...
, the
Laboratory of Molecular Biology The Medical Research Council (MRC) Laboratory of Molecular Biology (LMB) is a research institute in Cambridge, England, involved in the revolution in molecular biology which occurred in the 1950–60s. Since then it has remained a major medical ...
(and its precursors) at
Cambridge Cambridge ( ) is a university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cambridge becam ...
, and a handful of other institutions, the Pasteur Institute became a major center for molecular biology research in the late 1950s. Scientists at Cambridge, led by
Max Perutz Max Ferdinand Perutz (19 May 1914 – 6 February 2002) was an Austrian-born British molecular biologist, who shared the 1962 Nobel Prize for Chemistry with John Kendrew, for their studies of the structures of haemoglobin and myoglobin. He went ...
and
John Kendrew Sir John Cowdery Kendrew, (24 March 1917 – 23 August 1997) was an English biochemist, crystallographer, and science administrator. Kendrew shared the 1962 Nobel Prize in Chemistry with Max Perutz, for their work at the Cavendish Lab ...
, focused on the rapidly developing field of
structural biology Structural biology is a field that is many centuries old which, and as defined by the Journal of Structural Biology, deals with structural analysis of living material (formed, composed of, and/or maintained and refined by living cells) at every le ...
, combining X-ray crystallography with Molecular modelling and the new computational possibilities of digital computing (benefiting both directly and indirectly from the military funding of science). A number of biochemists led by
Frederick Sanger Frederick Sanger (; 13 August 1918 – 19 November 2013) was an English biochemist who received the Nobel Prize in Chemistry twice. He won the 1958 Chemistry Prize for determining the amino acid sequence of insulin and numerous other pr ...
later joined the Cambridge lab, bringing together the study of
macromolecular A macromolecule is a very large molecule important to biophysical processes, such as a protein or nucleic acid. It is composed of thousands of covalently bonded atoms. Many macromolecules are polymers of smaller molecules called monomers. The ...
structure and function. At the Pasteur Institute, François Jacob and Jacques Monod followed the 1959 PaJaMo experiment with a series of publications regarding the ''lac'' operon that established the concept of
gene regulation Regulation of gene expression, or gene regulation, includes a wide range of mechanisms that are used by cells to increase or decrease the production of specific gene products ( protein or RNA). Sophisticated programs of gene expression are w ...
and identified what came to be known as messenger RNA. By the mid-1960s, the intellectual core of molecular biology—a model for the molecular basis of metabolism and reproduction— was largely complete. The late 1950s to the early 1970s was a period of intense research and institutional expansion for molecular biology, which had only recently become a somewhat coherent discipline. In what organismic biologist E. O. Wilson called "The Molecular Wars", the methods and practitioners of molecular biology spread rapidly, often coming to dominate departments and even entire disciplines. Molecularization was particularly important in genetics,
immunology Immunology is a branch of medicineImmunology for Medical Students, Roderick Nairn, Matthew Helbert, Mosby, 2007 and biology that covers the medical study of immune systems in humans, animals, plants and sapient species. In such we can see ther ...
,
embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos an ...
, and
neurobiology Neuroscience is the scientific study of the nervous system (the brain, spinal cord, and peripheral nervous system), its functions and disorders. It is a multidisciplinary science that combines physiology, anatomy, molecular biology, developme ...
, while the idea that life is controlled by a " genetic program"—a metaphor Jacob and Monod introduced from the emerging fields of
cybernetics Cybernetics is a wide-ranging field concerned with circular causality, such as feedback, in regulatory and purposive systems. Cybernetics is named after an example of circular causal feedback, that of steering a ship, where the helmsperson ma ...
and computer science—became an influential perspective throughout biology. Immunology in particular became linked with molecular biology, with innovation flowing both ways: the clonal selection theory developed by
Niels Jerne Niels is a male given name, equivalent to Nicholas, which is common in Denmark, Belgium, Norway (formerly) and the Netherlands. The Norwegian and Swedish variant is Nils. The name is a developed short form of Nicholas or Greek Nicolaos after Sain ...
and
Frank Macfarlane Burnet Sir Frank Macfarlane Burnet, (3 September 1899 – 31 August 1985), usually known as Macfarlane or Mac Burnet, was an Australian virologist known for his contributions to immunology. He won a Nobel Prize in 1960 for predicting acquired immune ...
in the mid-1950s helped shed light on the general mechanisms of protein synthesis. Resistance to the growing influence of molecular biology was especially evident in evolutionary biology. Protein sequencing had great potential for the quantitative study of evolution (through the
molecular clock hypothesis The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleotid ...
), but leading evolutionary biologists questioned the relevance of molecular biology for answering the big questions of evolutionary causation. Departments and disciplines fractured as organismic biologists asserted their importance and independence:
Theodosius Dobzhansky Theodosius Grigorievich Dobzhansky (russian: Феодо́сий Григо́рьевич Добржа́нский; uk, Теодо́сій Григо́рович Добржа́нський; January 25, 1900 – December 18, 1975) was a prominent ...
made the famous statement that "
nothing in biology makes sense except in the light of evolution "Nothing in Biology Makes Sense Except in the Light of Evolution" is a 1973 essay by the evolutionary biologist Theodosius Dobzhansky, criticising anti-evolution creationism and espousing theistic evolution. The essay was first published in '' ...
" as a response to the molecular challenge. The issue became even more critical after 1968;
Motoo Kimura (November 13, 1924 – November 13, 1994) was a Japanese biologist best known for introducing the neutral theory of molecular evolution in 1968. He became one of the most influential theoretical population geneticists. He is remembered in genet ...
's
neutral theory of molecular evolution The neutral theory of molecular evolution holds that most evolutionary changes occur at the molecular level, and most of the variation within and between species are due to random genetic drift of mutant alleles that are selectively neutral. The ...
suggested that natural selection was not the ubiquitous cause of evolution, at least at the molecular level, and that molecular evolution might be a fundamentally different process from morphological evolution. (Resolving this "molecular/morphological paradox" has been a central focus of molecular evolution research since the 1960s.)


Biotechnology, genetic engineering, and genomics

Biotechnology Biotechnology is the integration of natural sciences and engineering sciences in order to achieve the application of organisms, cells, parts thereof and molecular analogues for products and services. The term ''biotechnology'' was first used by ...
in the general sense has been an important part of biology since the late 19th century. With the industrialization of
brewing Brewing is the production of beer by steeping a starch source (commonly cereal grains, the most popular of which is barley) in water and fermenting the resulting sweet liquid with yeast. It may be done in a brewery by a commercial brewer, ...
and
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
, chemists and biologists became aware of the great potential of human-controlled biological processes. In particular, fermentation proved a great boon to chemical industries. By the early 1970s, a wide range of biotechnologies were being developed, from drugs like penicillin and
steroids A steroid is a biologically active organic compound with four rings arranged in a specific molecular configuration. Steroids have two principal biological functions: as important components of cell membranes that alter membrane fluidity; and ...
to foods like ''
Chlorella ''Chlorella'' is a genus of about thirteen species of single-celled green algae belonging to the division Chlorophyta. The cells are spherical in shape, about 2 to 10 μm in diameter, and are without flagella. Their chloroplasts contain th ...
'' and single-cell protein to
gasohol Several common ethanol fuel mixtures are in use around the world. The use of pure hydrous or anhydrous ethanol in internal combustion engines (ICEs) is only possible if the engines are designed or modified for that purpose, and used only in autom ...
—as well as a wide range of
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two dif ...
high-yield crops and agricultural technologies, the basis for the
Green Revolution The Green Revolution, also known as the Third Agricultural Revolution, was a period of technology transfer initiatives that saw greatly increased crop yields and agricultural production. These changes in agriculture began in developed countrie ...
.


Recombinant DNA

Biotechnology in the modern sense of genetic engineering began in the 1970s, with the invention of
recombinant DNA Recombinant DNA (rDNA) molecules are DNA molecules formed by laboratory methods of genetic recombination (such as molecular cloning) that bring together genetic material from multiple sources, creating sequences that would not otherwise be fou ...
techniques. Restriction enzymes were discovered and characterized in the late 1960s, following on the heels of the isolation, then duplication, then synthesis of viral genes. Beginning with the lab of Paul Berg in 1972 (aided by '' EcoRI'' from
Herbert Boyer Herbert Wayne "Herb" Boyer (born July 10, 1936) is an American biotechnologist, researcher and entrepreneur in biotechnology. Along with Stanley N. Cohen and Paul Berg he discovered a method to coax bacteria into producing foreign proteins, the ...
's lab, building on work with ligase by
Arthur Kornberg Arthur Kornberg (March 3, 1918 – October 26, 2007) was an American biochemist who won the Nobel Prize in Physiology or Medicine 1959 for the discovery of "the mechanisms in the biological synthesis of ribonucleic acid and deoxyribonucleic aci ...
's lab), molecular biologists put these pieces together to produce the first transgenic organisms. Soon after, others began using plasmid vectors and adding genes for
antibiotic resistance Antimicrobial resistance (AMR) occurs when microbes evolve mechanisms that protect them from the effects of antimicrobials. All classes of microbes can evolve resistance. Fungi evolve antifungal resistance. Viruses evolve antiviral resistance. ...
, greatly increasing the reach of the recombinant techniques. Wary of the potential dangers (particularly the possibility of a prolific bacteria with a viral cancer-causing gene), the scientific community as well as a wide range of scientific outsiders reacted to these developments with both enthusiasm and fearful restraint. Prominent molecular biologists led by Berg suggested a temporary moratorium on recombinant DNA research until the dangers could be assessed and policies could be created. This moratorium was largely respected, until the participants in the 1975
Asilomar Conference on Recombinant DNA The Asilomar Conference on Recombinant DNA was an influential conference organized by Paul Berg, Maxine Singer, and colleagues to discuss the potential biohazards and regulation of biotechnology, held in February 1975 at a conference center at A ...
created policy recommendations and concluded that the technology could be used safely. Following Asilomar, new genetic engineering techniques and applications developed rapidly. DNA sequencing methods improved greatly (pioneered by
Frederick Sanger Frederick Sanger (; 13 August 1918 – 19 November 2013) was an English biochemist who received the Nobel Prize in Chemistry twice. He won the 1958 Chemistry Prize for determining the amino acid sequence of insulin and numerous other pr ...
and Walter Gilbert), as did oligonucleotide synthesis and transfection techniques. Researchers learned to control the expression of transgenes, and were soon racing—in both academic and industrial contexts—to create organisms capable of expressing human genes for the production of human hormones. However, this was a more daunting task than molecular biologists had expected; developments between 1977 and 1980 showed that, due to the phenomena of split genes and splicing, higher organisms had a much more complex system of gene expression than the bacteria models of earlier studies. The first such race, for synthesizing human insulin, was won by
Genentech Genentech, Inc., is an American biotechnology corporation headquartered in South San Francisco, California. It became an independent subsidiary of Roche in 2009. Genentech Research and Early Development operates as an independent center within ...
. This marked the beginning of the biotech boom (and with it, the era of
gene patent A biological patent is a patent on an invention in the field of biology that by law allows the patent holder to exclude others from making, using, selling, or importing the protected invention for a limited period of time. The scope and reach of ...
s), with an unprecedented level of overlap between biology, industry, and law.


Molecular systematics and genomics

By the 1980s, protein sequencing had already transformed methods of
scientific classification Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. A ...
of organisms (especially
cladistics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived cha ...
) but biologists soon began to use RNA and DNA sequences as
characters Character or Characters may refer to: Arts, entertainment, and media Literature * ''Character'' (novel), a 1936 Dutch novel by Ferdinand Bordewijk * ''Characters'' (Theophrastus), a classical Greek set of character sketches attributed to The ...
; this expanded the significance of molecular evolution within evolutionary biology, as the results of molecular systematics could be compared with traditional evolutionary trees based on
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
. Following the pioneering ideas of
Lynn Margulis Lynn Margulis (born Lynn Petra Alexander; March 5, 1938 – November 22, 2011) was an American evolutionary biologist, and was the primary modern proponent for the significance of symbiosis in evolution. Historian Jan Sapp has said that "Lynn Ma ...
on endosymbiotic theory, which holds that some of the
organelles In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
of eukaryotic cells originated from free living prokaryotic organisms through
symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasi ...
relationships, even the overall division of the tree of life was revised. Into the 1990s, the five domains (Plants, Animals, Fungi, Protists, and Monerans) became three (the Archaea, the Bacteria, and the Eukarya) based on
Carl Woese Carl Richard Woese (; July 15, 1928 – December 30, 2012) was an American microbiologist and biophysicist. Woese is famous for defining the Archaea (a new domain of life) in 1977 through a pioneering phylogenetic taxonomy of 16S ribosomal RNA, ...
's pioneering molecular systematics work with
16S rRNA 16S rRNA may refer to: * 16S ribosomal RNA 16 S ribosomal RNA (or 16 S rRNA) is the RNA component of the 30S subunit of a prokaryotic ribosome ( SSU rRNA). It binds to the Shine-Dalgarno sequence and provides most of the SSU structure. The g ...
sequencing. The development and popularization of the
polymerase chain reaction The polymerase chain reaction (PCR) is a method widely used to rapidly make millions to billions of copies (complete or partial) of a specific DNA sample, allowing scientists to take a very small sample of DNA and amplify it (or a part of it) ...
(PCR) in mid-1980s (by
Kary Mullis Kary Banks Mullis (December 28, 1944August 7, 2019) was an American biochemist. In recognition of his role in the invention of the polymerase chain reaction (PCR) technique, he shared the 1993 Nobel Prize in Chemistry with Michael Smith and was ...
and others at Cetus Corp.) marked another watershed in the history of modern biotechnology, greatly increasing the ease and speed of genetic analysis. Coupled with the use of expressed sequence tags, PCR led to the discovery of many more genes than could be found through traditional biochemical or genetic methods and opened the possibility of sequencing entire genomes. The unity of much of the morphogenesis of organisms from fertilized egg to adult began to be unraveled after the discovery of the
homeobox A homeobox is a DNA sequence, around 180 base pairs long, that regulates large-scale anatomical features in the early stages of embryonic development. For instance, mutations in a homeobox may change large-scale anatomical features of the full-g ...
genes, first in fruit flies, then in other insects and animals, including humans. These developments led to advances in the field of evolutionary developmental biology towards understanding how the various
body plan A body plan, ( ), or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually applied to animals, envisages a "blueprin ...
s of the animal phyla have evolved and how they are related to one another. The
Human Genome Project The Human Genome Project (HGP) was an international scientific research project with the goal of determining the base pairs that make up human DNA, and of identifying, mapping and sequencing all of the genes of the human genome from both a ph ...
—the largest, most costly single biological study ever undertaken—began in 1988 under the leadership of
James D. Watson James Dewey Watson (born April 6, 1928) is an American molecular biologist, geneticist, and zoologist. In 1953, he co-authored with Francis Crick the academic paper proposing the double helix structure of the DNA molecule. Watson, Crick and ...
, after preliminary work with genetically simpler model organisms such as '' E. coli'', '' S. cerevisiae'' and '' C. elegans''.
Shotgun sequencing In genetics, shotgun sequencing is a method used for sequencing random DNA strands. It is named by analogy with the rapidly expanding, quasi-random shot grouping of a shotgun. The chain-termination method of DNA sequencing ("Sanger sequencing ...
and gene discovery methods pioneered by Craig Venter—and fueled by the financial promise of gene patents with
Celera Genomics Celera is a subsidiary of Quest Diagnostics which focuses on genetic sequencing and related technologies. It was founded in 1998 as a business unit of Applera, spun off into an independent company in 2008, and finally acquired by Quest Diagnosti ...
— led to a public–private sequencing competition that ended in compromise with the first draft of the human DNA sequence announced in 2000.Davies, ''Cracking the Genome'', Introduction; see also: Sulston, ''The Common Thread''


Twenty-first century biological sciences

At the beginning of the 21st century, biological sciences converged with previously differentiated new and classic disciplines like Physics into research fields like
Biophysics Biophysics is an interdisciplinary science that applies approaches and methods traditionally used in physics to study biological phenomena. Biophysics covers all scales of biological organization, from molecular to organismic and populations. ...
. Advances were made in analytical chemistry and physics instrumentation including improved sensors, optics, tracers, instrumentation, signal processing, networks, robots, satellites, and compute power for data collection, storage, analysis, modeling, visualization, and simulations. These technological advances allowed theoretical and experimental research including internet publication of molecular biochemistry,
biological system A biological system is a complex network which connects several biologically relevant entities. Biological organization spans several scales and are determined based different structures depending on what the system is. Examples of biological syst ...
s, and ecosystems science. This enabled worldwide access to better measurements, theoretical models, complex simulations, theory predictive model experimentation, analysis, worldwide internet observational data reporting, open peer-review, collaboration, and internet publication. New fields of biological sciences research emerged including
Bioinformatics Bioinformatics () is an interdisciplinary field that develops methods and software tools for understanding biological data, in particular when the data sets are large and complex. As an interdisciplinary field of science, bioinformatics combine ...
, Neuroscience, Theoretical biology,
Computational genomics Computational genomics refers to the use of computational and statistical analysis to decipher biology from genome sequences and related data, including both DNA and RNA sequence as well as other "post-genomic" data (i.e., experimental data obtaine ...
,
Astrobiology Astrobiology, and the related field of exobiology, is an interdisciplinary scientific field that studies the origins, early evolution, distribution, and future of life in the universe. Astrobiology is the multidisciplinary field that investig ...
and Synthetic Biology.


See also

* History of botany * Outline of biology * Timeline of biology and organic chemistry


References


Citations


Sources

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Bowler, Peter J. Peter J. Bowler (born 8 October 1944) is a historian of biology who has written extensively on the history of evolutionary thought, the history of the environmental sciences, and on the history of genetics. His 1984 book, ''Evolution: The His ...
''The Earth Encompassed: A History of the Environmental Sciences''. W. W. Norton & Company: New York, 1992. *
Bowler, Peter J. Peter J. Bowler (born 8 October 1944) is a historian of biology who has written extensively on the history of evolutionary thought, the history of the environmental sciences, and on the history of genetics. His 1984 book, ''Evolution: The His ...
''The Eclipse of Darwinism: Anti-Darwinian Evolution Theories in the Decades around 1900''. The Johns Hopkins University Press: Baltimore, 1983. *
Bowler, Peter J. Peter J. Bowler (born 8 October 1944) is a historian of biology who has written extensively on the history of evolutionary thought, the history of the environmental sciences, and on the history of genetics. His 1984 book, ''Evolution: The His ...
''Evolution: The History of an Idea''. University of California Press, 2003. . * Browne, Janet. ''The Secular Ark: Studies in the History of Biogeography''. Yale University Press: New Haven, 1983. * Bud, Robert. ''The Uses of Life: A History of Biotechnology''. Cambridge University Press: London, 1993. * Caldwell, John. "Drug metabolism and pharmacogenetics: the British contribution to fields of international significance." ''British Journal of Pharmacology'', Vol. 147, Issue S1 (January 2006), pp S89–S99. * Coleman, William ''Biology in the Nineteenth Century: Problems of Form, Function, and Transformation''. Cambridge University Press: New York, 1977. * Creager, Angela N. H. ''The Life of a Virus: Tobacco Mosaic Virus as an Experimental Model, 1930–1965''. University of Chicago Press: Chicago, 2002. * Creager, Angela N. H. "Building Biology across the Atlantic," essay review in ''Journal of the History of Biology'', Vol. 36, No. 3 (September 2003), pp. 579–589. * de Chadarevian, Soraya. ''Designs for Life: Molecular Biology after World War II''. Cambridge University Press: Cambridge, 2002. * Dietrich, Michael R. "Paradox and Persuasion: Negotiating the Place of Molecular Evolution within Evolutionary Biology," in ''Journal of the History of Biology'', Vol. 31 (1998), pp. 85–111. * Davies, Kevin. ''Cracking the Genome: Inside the Race to Unlock Human DNA''. The Free Press: New York, 2001. * Fruton, Joseph S. ''Proteins, Enzymes, Genes: The Interplay of Chemistry and Biology''. Yale University Press: New Haven, 1999. * Gottweis, Herbert. ''Governing Molecules: The Discursive Politics of Genetic Engineering in Europe and the United States''. MIT Press: Cambridge, MA, 1998. * Gould, Stephen Jay. ''The Structure of Evolutionary Theory''. The Belknap Press of Harvard University Press: Cambridge, 2002. * Hagen, Joel B. ''An Entangled Bank: The Origins of Ecosystem Ecology''. Rutgers University Press: New Brunswick, 1992. * Hall, Stephen S. ''Invisible Frontiers: The Race to Synthesize a Human Gene''. Atlantic Monthly Press: New York, 1987. * Holmes, Frederic Lawrence. ''Meselson, Stahl, and the Replication of DNA: A History of "The Most Beautiful Experiment in Biology"''. Yale University Press: New Haven, 2001. * Junker, Thomas. ''Geschichte der Biologie''. C. H. Beck: München, 2004. * Kay, Lily E. ''The Molecular Vision of Life: Caltech, The Rockefeller Foundation, and the Rise of the New Biology''. Oxford University Press: New York, 1993. * Kohler, Robert E. ''Lords of the Fly: ''Drosophila'' Genetics and the Experimental Life''. Chicago University Press: Chicago, 1994. * Kohler, Robert E. ''Landscapes and Labscapes: Exploring the Lab-Field Border in Biology''. University of Chicago Press: Chicago, 2002. * Krimsky, Sheldon. ''Biotechnics and Society: The Rise of Industrial Genetics''. Praeger Publishers: New York, 1991. * Larson, Edward J. ''Evolution: The Remarkable History of a Scientific Theory''. The Modern Library: New York, 2004. * * Lovejoy, Arthur O. ''The Great Chain of Being: A Study of the History of an Idea''. Harvard University Press, 1936. Reprinted by Harper & Row, , 2005 paperback: . * Magner, Lois N. ''A History of the Life Sciences'', third edition. Marcel Dekker, Inc.: New York, 2002. * Mason, Stephen F. ''A History of the Sciences''. Collier Books: New York, 1956. * Mayr, Ernst. ''The Growth of Biological Thought: Diversity, Evolution, and Inheritance''. The Belknap Press of Harvard University Press: Cambridge, Massachusetts, 1982. * Mayr, Ernst and William B. Provine, eds. ''The Evolutionary Synthesis: Perspectives on the Unification of Biology''. Harvard University Press: Cambridge, 1998. * Morange, Michel. ''A History of Molecular Biology'', translated by Matthew Cobb. Harvard University Press: Cambridge, 1998. * Rabinbach, Anson. ''The Human Motor: Energy, Fatigue, and the Origins of Modernity''. University of California Press, 1992. * Rabinow, Paul. ''Making PCR: A Story of Biotechnology''. University of Chicago Press: Chicago, 1996. * Rudwick, Martin J.S. ''The Meaning of Fossils''. The University of Chicago Press: Chicago, 1972. * Raby, Peter. ''Bright Paradise: Victorian Scientific Travellers''. Princeton University Press: Princeton, 1997. * Rothman, Sheila M. and David J. Rothman. ''The Pursuit of Perfection: The Promise and Perils of Medical Enhancement''. Vintage Books: New York, 2003. * Sapp, Jan. ''Genesis: The Evolution of Biology''. Oxford University Press: New York, 2003. * Secord, James A. ''Victorian Sensation: The Extraordinary Publication, Reception, and Secret Authorship of ''Vestiges of the Natural History of Creation. University of Chicago Press: Chicago, 2000. * Serafini, Anthony ''The Epic History of Biology'', Perseus Publishing, 1993. * Sulston, John. ''The Common Thread: A Story of Science, Politics, Ethics and the Human Genome''. National Academy Press, 2002. * Smocovitis, Vassiliki Betty. ''Unifying Biology: The Evolutionary Synthesis and Evolutionary Biology''. Princeton University Press: Princeton, 1996. * Summers, William C. ''Félix d'Herelle and the Origins of Molecular Biology'', Yale University Press: New Haven, 1999. * Sturtevant, A. H.
A History of Genetics
'. Cold Spring Harbor Laboratory Press: Cold Spring Harbor, 2001. * Thackray, Arnold, ed. ''Private Science: Biotechnology and the Rise of the Molecular Sciences''. University of Pennsylvania Press: Philadelphia, 1998. * Wilson, Edward O. ''Naturalist''. Island Press, 1994. * Zimmer, Carl. ''Evolution: the triumph of an idea''. HarperCollins: New York, 2001.


External links


International Society for History, Philosophy, and Social Studies of Biology
– professional history of biology organization
History of Biology
– Historyworld article
History of Biology
at Bioexplorer.Net – a collection of history of biology links
Biology
– historically oriented article on Citizendium
Miall, L. C. (1911) History of biology.
Watts & Co. London * {{DEFAULTSORT:History Of Biology Articles containing video clips