The induced-fit theory

The key–lock hypothesis does not fully account for enzymatic action; i.e., certain properties of enzymes cannot be accounted for by the simple relationship between enzyme and substrate proposed by the key–lock hypothesis. A theory called the induced-fit theory retains the key–lock idea of a fit of the substrate at the active site but postulates in addition that the substrate must do more than simply fit into the already preformed shape of an active site. Rather, the theory states, the binding of the substrate to the enzyme must cause a change in the shape of the enzyme that results in the proper alignment of the catalytic groups on its surface. This concept has been likened to the fit of a hand in a glove, the hand (substrate) inducing a change in the shape of the glove (enzyme). Although some enzymes appear to function according to the older key–lock hypothesis, most apparently function according to the induced-fit theory.

Portion of the Encyclopædia Britannica Article

The String Theory

In particle physics, a theory that attempts to merge quantum mechanics with Albert Einstein's general theory of relativity. The name string theory comes from the modeling of subatomic particles as tiny one-dimensional “stringlike” entities rather than the more conventional approach in which they are modeled as zero-dimensional point particles. The theory envisions that a string undergoing a particular mode of vibration corresponds to a particle with definite properties such as mass and charge. In the 1980s, physicists realized that string theory had the potential to incorporate all four of nature's forces—gravity, electromagnetism, strong force, and weak force—and all types of matter in a single quantum mechanical framework, suggesting that it might be the long-sought unified field theory. While string theory is still a vibrant area of research that is undergoing rapid development, it remains a purely mathematical construct because it has yet to make contact with experimental observations.
Relativity and quantum mechanics

In 1905 Einstein unified space and time (see space-time) with his special theory of relativity, showing that motion through space affects the passage of time. In 1915 Einstein further unified space, time, and gravitation with his general theory of relativity, showing that warps and curves in space and time are responsible for the force of gravity. These were monumental achievements, but Einstein dreamed of an even grander unification. He envisioned one powerful framework that would account for space, time, and all of nature's forces—something he called a unified theory. For the last three decades of his life, Einstein relentlessly pursued his quixotic vision. Although from time to time rumours spread that he had succeeded, closer scrutiny always dashed such hopes. Most of Einstein's contemporaries considered the search for a unified theory to be a hopeless, if not misguided, quest.
In contrast, the primary concern of theoretical physicists from the 1920s onward was quantum mechanics—the emerging framework for describing atomic and subatomic processes. Particles at these scales have such tiny masses that gravity is essentially irrelevant in their interactions, and so for decades quantum mechanical calculations generally ignored relativistic effects. Instead, by the late 1960s the focus was on a different force—the strong force, which binds together the protons and neutrons within atomic nuclei. Gabriele Veneziano, a young theorist working at the European Organization for Nuclear Research (CERN), contributed a key breakthrough in 1968 with his realization that a 200-year-old formula, the Euler beta function, was capable of explaining much of the data on the strong force then being collected at various particle accelerators around the world. A few years later, three physicists—Leonard Susskind of Stanford University, Holger Nielsen of the Niels Bohr Institute, and Yoichiro Nambu of the University of Chicago—significantly amplified Veneziano's insight by showing that the mathematics underlying his proposal described the vibrational motion of minuscule filaments of energy that resemble tiny strands of string, inspiring the name string theory. Roughly speaking, the theory suggested that the strong force amounted to strings tethering together particles attached to the strings' endpoints.

Predictions and theoretical difficulties

String theory was an intuitively attractive proposal, but by the mid-1970s more-refined measurements of the strong force had deviated from its predictions, leading most researchers to conclude that string theory had no relevance to the physical universe, no matter how elegant the mathematical theory. Nevertheless, a small number of physicists continued to pursue string theory. In 1974 John Schwarz of the California Institute of Technology and Joel Scherk of the École Normale Supérieure and, independently, Tamiaki Yoneya of Hokkaido University came to a radical conclusion. They suggested that one of the supposedly failed predictions of string theory—the existence of a particular massless particle that no experiment studying the strong force had ever encountered—was actually evidence of the very unification Einstein had anticipated.
Although no one had succeeded in merging general relativity and quantum mechanics, preliminary work had established that such a union would require precisely the massless particle predicted by string theory. A few physicists argued that string theory, by having this particle built into its fundamental structure, had united the laws of the large (general relativity) and the laws of the small (quantum mechanics). Rather than merely being a description of the strong force, these physicists contended, string theory required reinterpretation as a critical step toward Einstein's unified theory.
The announcement was universally ignored. String theory had already failed in its first incarnation as a description of the strong force, and many felt it was unlikely that it would now prevail as the solution to an even more difficult problem. This view was bolstered by string theory's suffering from its own theoretical problems. For one, some of its equations showed signs of being inconsistent; for another, the mathematics of the theory demanded the universe have not just the three spatial dimensions of common experience but six others (for a total of nine spatial dimensions, or a total of ten space-time dimensions).

Dimensions and vibrations

Because of these obstacles, the number of physicists working on the theory had dropped to two—Schwarz and Michael Green, of Queen Mary College, London—by the mid-1980s. But in 1984 these two die-hard string theorists achieved a major breakthrough. Through a remarkable calculation, they proved that the equations of string theory were consistent after all. By the time word of this result had spread throughout the physics community, hundreds of researchers had dropped what they were working on and turned their full attention to string theory.
Within a few months, string theory's unified framework took shape. Much as different vibrational patterns of a violin string play different musical notes, the different vibrations of the tiny strands in string theory were imagined to yield different particles of nature. According to the theory, the strings are so small that they appear to be points—as particles had long been thought to be—but in reality they have length (about 10-33 cm); the mass and charge of a particle is determined by how a string vibrates. For example, string theory posits that an electron is a string undergoing one particular vibrational pattern; a quark is imagined as a string undergoing a different vibrational pattern. Crucially, among the vibrational patterns, physicists argued, would also be the particles found by experiment to communicate nature's forces. Thus, string theory was proposed as the sought-for unification of all forces and all matter.
What of the six extra spatial dimensions required by string theory? Following a suggestion made in the 1920s by Theodor Kaluza of Germany and Oskar Klein of Sweden, string theorists envisioned that dimensions come in two distinct varieties. Like the unfurled length of a long garden hose, dimensions can be big and easy to see. But like the shorter, circular girth of the garden hose, dimensions can also be far smaller and more difficult to detect. This becomes more apparent by imagining that the circular cross section of the garden hose is shrunk ever smaller, below what can be seen with the naked eye, thereby misleading a casual observer into thinking the garden hose has only one dimension, its length. Similarly, according to string theory, the three dimensions of common experience are large and hence manifest, while the other six dimensions are crumpled so small that they have so far evaded detection.
During the decade from 1984 to 1994, many theoretical physicists strove to fulfill string theory's promise by developing this abstract, wholly mathematical framework into a concrete, predictive theory of nature. Because the infinitesimal size of strings has precluded their direct detection, theorists have sought to extract indirect implications of the theory that might be testable. In this regard, the extra dimensions of string theory have proved a major hurdle. Imagining these extra dimensions as small and hidden is a reasonable explanation for their apparent absence. However, also because strings are so small, they would vibrate in every dimension, not just in the usual three dimensions. Studies showed that, much as the shape and size of a French horn affect the vibrational patterns of airstreams coursing through the instrument, the exact shape and size of the extra dimensions would affect how strings vibrate. And since the strings' vibrations determine quantities such as particle masses and charges, predictivity requires knowledge of the geometrical form of the extra dimensions. Unfortunately, the equations of string theory allow the extra dimensions to take many different geometric forms, making it difficult to extract definitive testable predictions.

M-theory

By the mid-1990s, these and other obstacles were again eroding the ranks of string theorists. But in 1995 another breakthrough reinvigorated the field. Edward Witten of the Institute for Advanced Study, building on contributions of many other physicists, proposed a new set of techniques that refined the approximate equations on which all work in string theory had so far been based. These techniques helped reveal a number of new features of string theory. Most dramatically, these more exact equations showed that string theory has not six but seven extra spatial dimensions; the more exact equations also revealed ingredients in string theory besides strings—membranelike objects of various dimensions, collectively called branes. Finally, the new techniques established that various versions of string theory developed over the preceding decades were essentially all the same. Theorists call this unification of formerly distinct string theories by a new name, M-theory, with the meaning of M being deferred until the theory is more fully understood.
Today, the understanding of many facets of string theory is still in its formative stage. Researchers recognize that, although remarkable progress has been made over the past three decades, collectively the work is burdened by its piecemeal development, with incremental discoveries having been joined like pieces of a jigsaw puzzle. That the pieces fit coherently is impressive, but the larger picture they are filling out—the fundamental principle underlying the theory—remains mysterious. Equally pressing, the theory has yet to be supported by observations and hence remains a totally theoretical construct.
In the next decade this could change. An intriguing outcome of theoretical developments since 1995 is the recognition that strings and the extra dimensions might be significantly larger than previously thought. Rather than being 10-33 cm, studies with the more refined M-theory framework have established that strings could be larger by many orders of magnitude. If so, the next generation of particle accelerators (such as the Large Hadron Collider at CERN) may have enough energy to probe the physical properties of strings directly, providing the long-sought experimental confirmation of the theory.

Supersymmetry and cosmological signatures

The experiments at CERN will also search for evidence of supersymmetry, a mathematical property discovered within string theory that requires every known particle species to have a partner particle species, called superpartners. (This property accounts for string theory often being referred to as superstring theory.) As yet, no superpartner particles have been detected, but researchers believe this may be due to their weight—they are heavier than their known counterparts and require a machine at least as powerful as the Large Hadron Collider to produce them. If the superpartner particles are found, string theory still will not be proved correct, because more-conventional point-particle theories have also successfully incorporated supersymmetry into their mathematical structure. However, the discovery of supersymmetry would confirm an essential element of string theory and give circumstantial evidence that this approach to unification is on the right track.
Even if these accelerator-based tests are inconclusive, there is another way that string theory may one day be tested. Through its impact on the earliest, most extreme moments of the universe, the physics of string theory may have left faint cosmological signatures—for example, in the form of gravitational waves or a particular pattern of temperature variations in the cosmic microwave background radiation—that may be observable by the next generation of precision satellite-borne telescopes and detectors. It would be a fitting conclusion to Einstein's quest for unification if a theory of the smallest microscopic component of matter were confirmed through observations of the largest astronomical realms of the cosmos.

- Encyclopædia Britannica Article

The Cell Theory

Although the microscopists of the 17th century had made detailed descriptions of plant and animal structure and though Hooke had coined the term cell for the compartments he had observed in cork tissue, their observations lacked an underlying theoretical unity. It was not until 1838 that Matthias J. Schleiden, a German botanist interested in plant anatomy, stated, “the lower plants all consist of one cell, while the higher ones are composed of (many) individual cells.” When Schleiden's friend, the German physiologist Theodor Schwann, extended the cellular theory to include animals, he thereby brought about a rapprochement between botany and zoology. The formation of the cell theory—all plants and animals are made up of cells—marked a great conceptual advance in biology, and it resulted in renewed attention to the living processes that go on in cells.
In 1846, after several investigators had described the streaming movement of the cytoplasm in plant cells, Hugo von Mohl, a German botanist, coined the word protoplasm to designate the living substance of the cell. The concept of protoplasm as the physical basis of life led to the development of cell physiology.
A further extension of the cell theory was the development of cellular pathology by Rudolf Virchow, who established the relationship between abnormal events in the body and unusual cellular activities. This gave a new direction to the study of pathology and resulted in advances in medicine.
The detailed description of cell division was contributed by Eduard Strasburger, a German botanist, who observed the mitotic process in plant cells and further demonstrated that nuclei arise only from preexisting nuclei. The parallel work in mammals was done by the German anatomist Walther Flemming, who published his most important findings in Zellsubstanz, Kern und Zelltheilung (“Cell Substance, Nucleus and Cell Division”) in 1882.



The history of cell theory is a history of the actual observation of cells, because early prediction and speculation about the nature of the cell were generally unsuccessful. The decisive event that allowed the observation of cells was the invention of the microscope in the 17th century, after which interest in the invisible world was stimulated. Robert Hooke, who described cork and other plant tissues in 1665, introduced the term cell because the cellulose walls of dead cork cells reminded him of the blocks of cells occupied by monks. Even after the publication in 1672 of excellent pictures of plant tissues, no significance was attached to the contents within the cell walls. The magnifying powers of the microscope and the inadequacy of techniques for preparing cells for observation precluded a study of the intimate details of the cell contents. The inspired amateur of early microscopy Antonie van Leeuwenhoek, beginning in 1673, discovered blood cells, spermatozoa, and a lively world of “animalcules.” A new world of unicellular organisms was opened up. Such discoveries extended the known variety of living things but did not bring insight into their basic uniformity. Moreover, when Leeuwenhoek observed the swarming of his animalcules but failed to observe their division, he could only reinforce the idea that they arose spontaneously.
Cell theory was not formulated for nearly 200 years after the introduction of microscopy. Explanations for this delay range from the poor quality of the microscopes to the persistence of ancient ideas concerning the definition of a fundamental living unit. Many observations of cells were made, but apparently none of the observers was able to assert forcefully that cells were the units of biologic structure and function.
Three critical discoveries made during the 1830s, when improved microscopes with suitable lenses, higher powers of magnification without aberration, and more satisfactory illumination became available, were decisive events in the early development of cell theory. First, the nucleus was observed by Robert Brown in 1833 as a constant component of plant cells. Next, nuclei were also observed and recognized as such in some animal cells. Finally, a living substance called protoplasm was recognized within cells, its vitality made evident by its active streaming, or flowing, movements, especially in plant cells. After these three discoveries, cells, previously considered as mere pores in plant tissue, could no longer be thought of as empty, because they contained living material.
Two German biologists, Theodore Schwann and Matthias Schleiden, clearly stated in 1839 that cells are the “elementary particles of organisms” in both plants and animals and recognized that some organisms are unicellular and others multicellular. This statement was made in Schwann's Mikroskopische Untersuchungen über die Übereinstimmung in der Struktur und dem Wachstume der Tiere und Pflanzen (1839; Microscopical Researches into the Accordance in the Structure and Growth of Animals and Plants). Schleiden's contributions on plants were acknowledged by Schwann as the basis for his comparison of animal and plant structure.
Schleiden and Schwann's descriptive statements concerning the cellular basis of biologic structure are straightforward and acceptable to modern thought. They recognized the common features of cells to be membrane, nucleus, and cell body and described them in comparisons of various animal and plant tissues. A statement by Schleiden pointed toward the future direction of cell studies:
Each cell leads a double life: an independent one, pertaining to its own development alone; and another incidental, insofar as it has become an integral part of a plant. It is, however, easy to perceive that the vital process of the individual cells must form the first, absolutely indispensable fundamental basis, both as regards vegetable physiology and comparative physiology in general.
Schwann and Schleiden were not alone in contributing to this great generalization of natural science, for strong intimations of the cell theory occur in the work of their predecessors. Recognizing that the basic problem was the origin of cells, these early investigators invented a hypothesis of “free cell formation,” according to which cells developed de novo out of an unformed substance, a “cytoblastema,” by a sequence of events in which first the nucleolus develops, followed by the nucleus, the cell body, and finally the cell membrane. The best physical model of the generation of formed bodies then available was crystallization, and their theory was inspired by that model. In retrospect, the hypothesis of free cell formation would not seem to have been justified, however, since cell division, a feature not characteristic of crystallization processes, had frequently been observed by earlier microscopists, especially among single-celled organisms. Even though cell division was observed repeatedly in the following decades, the theory of free cell formation lingered throughout most of the 19th century; however, it came to be thought of more and more as a possible exception to the general principle of the reproduction of cells by division. The correct general principle was affirmed in 1855 by a German biologist of great prestige, Rudolph Virchow, who asserted that “omnis cellula e cellula” (“all cells come from cells”).

- Encyclopædia Britannica Article