In 1873 Italy, Camillo Golgi created the black reaction technique, which enabled scientists to stain and view the structure of neurons, the specialized cells that compose the nervous system. During the nineteenth century, scientists were studying cells and proposed cell theory, which describes the basic characteristics of cells as fundamental units of life. However, cell theory struggled to explain neurons as they are specialized cells and more complex in structure than cells of other tissues. Prior to Golgi’s black reaction, other neuron staining techniques did not enable scientists to clearly and completely view entire neurons without damaging the tissue and obscuring the form. By enabling scientists to study individual neurons and neural tissues, Golgi’s black reaction enables researchers to better study the nervous system and how it develops.

Santiago Felipe Ramon y Cajal investigated brains in the nineteenth and twentieth centuries in Spain. He identified and individuated many components of the brain, including the neuron and the axon. He used chick embryos instead of adult animals, then customary in brain research, to study the development and physiology of the cerebellum, spinal cord, and retina. Ramon y Cajal received the Nobel Prize in Physiology and Medicine in 1906, along with Camillo Golgi, for his work on the structure of the nervous system.

In 2007, Françoise Baylis and Jason Scott Robert published “Part-Human Chimeras: Worrying the Facts, Probing the Ethics” in The American Journal of Bioethics. Within their article, hereafter “Part-Human Chimeras,” the authors offer corrections on “Thinking About the Human Neuron Mouse,” a report published in The American Journal of Bioethics in 2007 by Henry Greely, Mildred K. Cho, Linda F. Hogle, and Debra M. Satz, which discussed the debate on the ethics of creating part-human chimeras. Chimeras are organisms that contain two or more genetically distinct cell lines. Both publications discuss chimeras with DNA from different species, specifically in response to studies in which scientists injected human brain cells into mice. “Part-Human Chimeras,” contributes to a chain of ethical and scientific discussion that occurred in the mid-2000s on whether people should be able to conduct research on chimeras, especially in embryos.

In 2011, Sonja Vernes and Simon Fisher performed a series of experiments to determine which developmental processes are controlled by the mouse protein Foxp2. Previous research showed that altering the Foxp2 protein changed how neurons grew, so Vernes and Fisher hypothesized that Foxp2 would affect gene networks that involved in the development of neurons, or nerve cells. Their results confirmed that Foxp2 affected the development of gene networks involved in the growth of neurons, as well as networks that are involved in cell specialization and cell communication. The researchers determined that Foxp2 is important for a variety of developmental processes such as motor control, language acquisition, and cognition.

Scientists use cerebral organoids, which are artificially produced miniature organs that represent embryonic or fetal brains and have many properties similar to them, to help them study developmental disorders like microcephaly. In human embryos, cerebral tissue in the form of neuroectoderm appears within the first nine weeks of human development, and it gives rise to the brain and spinal cord. In the twenty-first century, Juergen Knoblich and Madeleine Lancaster at the Institute of Molecular Biotechnology in Vienna, Austria, grew cerebral organoids from pluripotent stem cells as a model to study developmental disorders in embryonic and fetal brains. One such disorder is microcephaly, a condition in which brain size and the number of neurons in the brain are abnormally small. Scientists use cerebral organoids, which they've grown in labs, because they provide a manipulable model for studying how neural cells migrate during development, the timing of neural development, and how genetic errors can result in developmental disorders.

Apoptosis, or programmed cell death, is a mechanism in embryonic development that occurs naturally in organisms. Apoptosis is a different process from cell necrosis, which is uncontrolled cell death usually after infection or specific trauma. As cells rapidly proliferate during development, some of them undergo apoptosis, which is necessary for many stages in development, including neural development, reduction in egg cells (oocytes) at birth, as well as the shaping of fingers and vestigial organs in humans and other animals. Sydney Brenner, H. Robert Horvitz, and John E. Sulston received the Nobel Prize in Physiology or Medicine in 2002 for their work on the genetic regulation of organ development and programmed cell death. Research on cell lineages before and after embryonic development may lead to new ways to reduce or promote cell death, which can be important in preventing diseases such as Alzheimer's or cancer.

The neuron doctrine is a concept formed during the turn of the twentieth century that describes the properties of neurons, the specialized cells that compose the nervous system. The neuron doctrine was one of two major theories on the composition of the nervous system at the time. Advocates of the neuron doctrine claimed that the nervous system was composed of discrete cellular units. Proponents of the alternative reticular theory, on the other hand, argued that the entire nervous system was a continuous network of cells, without gaps or synapses between the cells. In 1873, physician and reticular theory supporter Camillo Golgi developed a staining technique called the black reaction, a neuron staining technique that allowed for complete visibility of nerve cells, which enabled scientists to view a complete neuron cell and its cellular structures. Later, neuroscientist Santiago Ramón y Cajal used the black reaction to show the existence of synapses, or gaps between neurons, and argued that his evidence supported the neuron doctrine. The confirmation of the neuron doctrine showed that neurons function as discrete and independent cells, not as a single network, within the nervous system.

In the nineteenth century, reticular theory aimed to describe the properties of neurons, the specialized cells which make up the nervous system, but was later disconfirmed by evidence. Reticular theory stated that the nervous system was composed of a continuous network of specialized cells without gaps (synapses), and was first proposed by researcher Joseph von Gerlach in Germany in 1871. Reticular theory played a significant role in developmental neurobiology as it enabled scientists to theorize how the form of neural cells functioned in the context of the broader nervous system, and although disproven, reticular theory contributed to the foundation of the neuron doctrine that informed the modern field of neurobiology.

Camillo Golgi studied the central nervous system during the late nineteenth and early twentieth centuries in Italy, and he developed a staining technique to visualize brain cells. Called the black reaction, Golgi’s staining technique enabled him to see the cellular structure of brain cells, called neurons, with much greater precision. Golgi also used the black reaction to identify structures within animal cells like the internal reticular apparatus that stores, packs, and modifies proteins, later named the Golgi apparatus in his honor. Golgi, along with Santiago Ramón y Cajal, received the Nobel Peace Prize in 1906 for their independent work on the structure of the nervous system. Golgi’s discovery of the black reaction enabled other scientists to better study the structure of the nervous system and its development.

The Golgi staining technique, also called the black reaction after the stain's color, was developed in the 1870s and 1880s in Italy to make brain cells (neurons) visible under the microscope. Camillo Golgi developed the technique while working with nervous tissue, which required Golgi to examine cell structure under the microscope. Golgi improved upon existing methods of staining, enabling scientists to view entire neurons for the first time and changing the way people discussed the development and composition of the brain's cells. Into the twenty-fist century, Golgi's staining method continued to inform research on the nervous system, particularly regarding embryonic development.

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