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After the first observations of life under the microscope, it took two centuries of research before
the ‘cell theory’, the idea that all living things are composed of cells or their products, was
formulated. It proved even harder to accept that individual cells also make up nervous tissue.
ith the invention of the microscope hoek (1632–1723). The particles that he saw for naturalists. He described the micro-
NATURE CELL BIOLOGY | VOL 1 | MAY 1999 | | cellbio.nature.com © 1999 Macmillan Magazines Ltd E13
The cell theory material to become a new cell. This theory Within the nucleus, the nucleolus and a
Hints at the idea that the cell is the basic of ‘free cell formation’ was reminiscent of stainable substance could be seen. Moreo-
component of living organisms emerged the old ‘spontaneous generation’ doctrine ver, a number of structures (ribbons, bands
well before 1838–39, which was when the (although as an intracellular variant), but and threads) appeared during cell division.
cell theory was officially formulated. Cells was refuted in the 1850s by Robert Remak As these structures could be heavily stained,
were not seen as undifferentiated struc- (1815–1865), Rudolf Virchow (1821–1902) they were called ‘‘chromatin’’ by Walther
tures. Some cellular components, such as and Albert Kölliker (1817–1905) who Flemming (1843–1905), who also intro-
the nucleus, had been visualized, and the showed that cells are formed through scis- duced the term ‘‘mitosis’’ in 1882 and gave
occurrence of these structures in cells of dif- sion of pre-existing cells7. Virchow’s apho- a superb description of its various
ferent tissues and organisms hinted at the rism omnis cellula e cellula (every cell from a processes18. Flemming observed the longi-
possibility that cells of similar organization pre-existing cell) thus became the basis of tudinal splitting of salamander chromo-
might underlie all living matter. the theory of tissue formation, even if the somes (a term introduced only in 1888 by
The abbot Felice Fontana (1730–1805) mechanisms of nuclear division were not Wilhelm Waldeyer, 1836–1921) during
glimpsed the nucleus in epithelial cells in understood at the time. metaphase and established that each half-
1781, but this structure had probably been Cell theory stimulated a reductionistic chromosome moves to the opposite pole of
observed in animal and plant cells in the approach to biological problems and the mitotic nucleus18. This process was also
first decades of the eighteenth century7,10. became the most general structural para- observed in plants, providing further evi-
The Scottish botanist Robert Brown (1773– digm in biology. It emphasized the concept dence of the deep unity of the living world.
1858) was the first to recognize the nucleus of the unity of life and brought about the
(a term that he introduced) as an essential concept of organisms as “republics of living The neuron theory
constituent of living cells (1831). In the elementary units”7. There was, however, a tissue that seemed to
leaves of orchids Brown observed “a single As well as being the fundamental unit of belie cell theory — nervous tissue. Because
circular areola, generally somewhat more life, the cell was also seen as the basic ele- of its softness and fragility, it was difficult to
opake than the membrane of the cell... This ment of pathological processes. Diseases handle and susceptible to deterioration. But
areola, or nucleus of the cell as perhaps it came to be considered (irrespective of the it was its structural complexity that pre-
might be termed, is not confined to the epi- causative agent) as an alteration of cells in vented a simple reduction to models
dermis, being also found not only in the the organism. Virchow’s Cellularpathologie derived from the cell theory. Nerve-cell
pubescence of the surface... but in many was the most important pathogenic concept bodies, nervous prolongations and nervous
cases in the parenchyma or internal cells of until, in this century, the theory of molecu- fibres were observed in the first half of the
the tissue”11. Brown recognized the general lar pathology was developed. nineteenth century. However, attempts at
occurrence of the nucleus in these cells and reconstructing a three-dimensional struc-
apparently thought of the organization of Protoplasmic constituents ture of the nervous system were frustrated
the plant in terms of cellular constituents. After Schleiden and Swann’s formulation of by the impossibility of determining the
Meanwhile, technical improvements in cell theory, the basic constituents of the cell exact relationships between cell bodies
microscopy were being made. The principal were considered to be a wall or a simple (somas), neuronal protoplasmic processes
drawback of microscopes since van Leeu- membrane, a viscous substance called ‘‘pro- (dendrites) and nervous fibres.
wenhoek’s time was what we now call ‘chro- toplasm’’ (a name now replaced by Köl- A book by Karl Deiters (1834–1863),
matic aberration’, which diminishes the liker’s term ‘‘cytoplasm’’), and the nucleus. published posthumously in 1865, contains
resolution power of the instrument at high It soon became evident that the protoplasm beautiful descriptions and drawings of
magnifications. Only in the 1830s were ach- was not a homogeneous fluid. Some biolo- nerve cells studied by using histological
romatic microscopes introduced, allowing gists regarded its fine structure as fibrillary, methods and microdissections made with
more precise histological observations. whereas others described a reticular, alveo- thin needles under the microscope (see
Improvements were also made in tissue- lar or granular protoplasmic architecture. photographs on next page)19. Deiters’s
preservation and -treating techniques. This discrepancy resulted partly from arte- nerve cells were characterized by a soma,
In 1838, the botanist Matthias Jakob factual and illusory images attributable to dendrites and a nerve prolongation (axon)
Schleiden (1804–1881) suggested that fixation and staining procedures that which showed no branching. Kölliker, in
every structural element of plants is com- caused a non-homogeneous precipitation the fifth edition of his important book on
posed of cells or their products12. The fol- of colloidal complexes. histology, published in 1867, proposed that
lowing year, a similar conclusion was However, some staining of real cellular sensory and motor cells of the right and left
elaborated for animals by the zoologist components led to the description of differ- halves of the spinal cord were linked “by
Theodor Schwann (1810–1882). He stated entiated elements, which were subsequently anastomoses” (direct fusion)20.
that “the elementary parts of all tissues are identified. The introduction of the oil- In 1872, the German histologist Joseph
formed of cells” and that “there is one uni- immersion lens in 1870, the development of Gerlach (1820–1896) expanded Kölliker’s
versal principle of development for the ele- the microtome technique and the use of view and proposed that, in all of the central
mentary parts of organisms... and this new fixing methods and dyes greatly nervous system, nerve cells established
principle is in the formation of cells”13. The improved microscopy. Towards the end of anastomoses with each other through a net-
conclusions of Schleiden and Schwann are the nineteenth century, the principal work formed by the minute branching of
considered to represent the official formu- organelles that are now considered to be their dendrites. According to this concept,
lation of ‘cell theory’ and their names are parts of the cell were identified. The term the network or reticulum was an essential
almost as closely linked to cell theory as are ‘‘ergastoplasm’’ (endoplasmic reticulum) element of grey matter that provided a sys-
those of Watson and Crick with the struc- was introduced in 1897 (ref. 15); mitochon- tem for anatomical and functional commu-
ture of DNA4,14. dria were observed by several authors and nications, a protoplasmic continuum from
According to Schleiden, however, the named by Carl Benda (1857–1933) in 1898 which nerve fibres originated21.
first phase of the generation of cells was the (ref. 16), the same year in which Camillo The most important breakthrough in
formation of a nucleus of ‘‘crystallization’’ Golgi (1843–1926) discovered the intracel- neurocytology and neuroanatomy came in
within the intracellular substance (which he lular apparatus that bears his name17. 1873 when Golgi developed the ‘black reac-
called the ‘‘cytoblast’’), with subsequent The protoplasm was not the only struc- tion’22, which he announced to a friend with
progressive enlargement of such condensed ture to have a heterogeneous appearance. these few words, “I am delighted that I have
E14 © 1999 Macmillan Magazines Ltd NATURE CELL BIOLOGY | VOL 1 | MAY 1999 | cellbio.nature.com
NATURE CELL BIOLOGY | VOL 1 | MAY 1999 | | cellbio.nature.com © 1999 Macmillan Magazines Ltd E15