Telomeres are sequences of DNA on the ends of chromosomes that protect chromosomes from sticking to each other or tangling, which could cause irregularities in normal DNA functions. As cells replicate, telomeres shorten at the end of chromosomes, which correlates to senescence or cellular aging. Integral to this process is telomerase, which is an enzyme that repairs telomeres and is present in various cells in the human body, especially during human growth and development. Telomeres and telomerase are required for normal human embryonic development because they protect DNA as it completes multiple rounds of replication.

Friedrich Leopold August Weismann published Das
Keimplasma: eine Theorie der Vererbung (The Germ-Plasm: a
Theory of Heredity, hereafter The Germ-Plasm) while
working at the University of Freiburg in Freiburg, Germany in 1892.
William N. Parker, a professor in the University College of South
Wales and Monmouthshire in Cardiff, UK, translated The
Germ-Plasm into English in 1893. In The Germ-Plasm,
Weismann proposed a theory of heredity based on the concept of the
germ plasm, a substance in the germ cell that carries hereditary information. The
Germ-Plasm compiled Weismann's theoretical work and analyses of
other biologists' experimental work in the 1880s, and it provided a
framework to study development, evolution and heredity. Weismann
anticipated that the germ-plasm theory would enable researchers to
investigate the functions and material of hereditary substances.

Barbara McClintock conducted experiments on corn (Zea mays) in the United States in the mid-twentieth century to study the structure and function of the chromosomes in the cells. McClintock researched how genes combined in corn and proposed mechanisms for how those interactions are regulated. McClintock received the Nobel Prize in Physiology or Medicine in 1983, the first woman to win the prize without sharing it. McClintock won the award for her introduction of the concept of transposons, also called jumping genes. McClintock conceptualized some genetic material as not static in structure and order, but as subject to re-arrangement and may be altered during development.

The Hayflick Limit is a concept that helps to explain the
mechanisms behind cellular aging. The concept states that a normal human
cell can only replicate and divide forty to sixty times before it
cannot divide anymore, and will break down by programmed cell death
or apoptosis. The concept of the Hayflick Limit revised Alexis
Carrel's earlier theory, which stated that cells can replicate
themselves infinitely. Leonard Hayflick developed the concept while
at the Wistar Institute in Philadelphia,
Pennsylvania, in 1965. In his 1974 book Intrinsic
Mutagenesis, Frank Macfarlane Burnet named the concept after
Hayflick. The concept of the Hayflick Limit helped scientists study
the effects of cellular aging on human populations from embryonic
development to death, including the discovery of the effects of
shortening repetitive sequences of DNA, called telomeres, on the
ends of chromosomes. Elizabeth Blackburn, Jack Szostak and Carol
Greider received the Nobel Prize in Physiology or Medicine in 2009
for their work on genetic structures related to the Hayflick
Limit.

Theophilus Shickel Painter studied the structure and
function of chromosomes in the US during in the early to mid-twentieth century. Painter worked at
the University of Texas at Austin in Austin, Texas. In the 1920s
and 1930s, Painter studied the chromosomes of the salivary gland
giant chromosomes of the fruit fly (Drosophila
melanogaster), with Hermann J. Muller. Muller and Painter
studied the ability of X-rays to cause changes in the chromosomes
of fruit flies. Painter also studied chromosomes in mammals.
He investigated the development of the male gamete, a process
called spermatogenesis, in several invertebrates and vertebrates,
including mammals. In addition, Painter studied the role the
Y-chromosome plays in the determination and development of the male
embryo. Painter's research concluded that egg cells fertilized by
sperm cell bearing an X-chromosome resulted in a female embryo,
whereas egg cells fertilized by a sperm cell carrying a
Y-chromosome resulted in a male embryo. Painter's work with
chromosomes helped other researchers determine that X- and
Y-chromosomes are responsible for sex determination.

Ontogeny and Phylogeny is a book published in 1977, in which the author Stephen J. Gould, who worked in the US, tells a history of the theory of recapitulation. A theory of recapitulation aims to explain the relationship between the embryonic development of an organism (ontogeny) and the evolution of that organism's species (phylogeny). Although there are several variations of recapitulationist theories, most claim that during embryonic development an organism repeats the adult stages of organisms from those species in it's evolutionary history. Gould suggests that, although fewer biologists invoked recapitulation theories in the twentieth century compared to those in the nineteenth and eighteenth centuries, some aspects of the theory of recapitulation remained important for understanding evolution. Gould notes that the concepts of acceleration and retardation during development entail that changes in developmental timing (heterochrony) can result in a trait appearing either earlier or later than normal in developmental processes. Gould argues that these changes in the timing of embryonic development provide the raw materials or novelties upon which natural selection acts.

The Spandrels of San Marco and the Panglossian Paradigm:
A Critique of the Adaptationist Programme, hereafter called
The Spandrels, is an article written by Stephen J. Gould and
Richard C. Lewontin published in the Proceedings of the Royal
Society of London in 1979. The paper emphasizes issues with
what the two authors call adaptationism or the adaptationist
programme as a framework to explain how species and traits evolved. The paper
is one in a series of works in which Gould emphasized the
role of development in evolutionary theories. The article suggests
that constraints on how organisms can develop and constraints on how species can evolve from others play a
central role in explaining the how species and traits evolve. The
authors note that organisms from different species develop as
embryos through stages similar across species, genera, and higher
classes. Gould and Lewontin hypothesize that those stages
constrained the possible pathways of evolution and has therefore
guided the history of life. Throughout the paper, the authors rely on analogy of some parts of organisms to architectural structures called spandrels, marked in this image as 'a'."

The Cell-Theory was written by Thomas Henry Huxley in Britain and published in 1853 by The British and Foreign Medico-Chirurgical Review. The twenty-two page article reviews twelve works on cell theory, including those in Germany by Caspar Friedrich Wolff in the eighteenth century and by Karl Ernst von Baer in the nineteenth century. Huxley spends much of The Cell-Theory on a cell theory proposed in the late 1830s by Matthias Schleiden and Theodor Schwann in Germany. Schleiden and Schwann maintained that the cell was the most fundamental unit of life and that the nucleus was the most significant cellular component. Huxley, instead, promoted an epigenetic theory of the cell, for which properties of life emerge from the outer cytoplasm, cell membrane, and wall (the periplast), as opposed to the inner contents of the cell, including the nucleus (the endoplast). Huxley's arguments in The Cell-Theory influenced future scientists about the role of epigenetic processes in embryology and development.

Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Organs is the thirteenth chapter of Charles Darwin's book The Origin of Species, first published in England in 1859. The book details part of Darwin's argument for the common ancestry of life and natural selection as the cause of speciation. In this chapter, Darwin summarizes the evidence for evolution by connecting observations of development in organisms to the processes of natural selection. Darwin shows how the theory of special creation, which claims that God directly created all organisms in their current form, is inferior to the theory of natural selection for its ability to explain the diversity of life. In this chapter, Darwin also discusses classification and homology as they relate to natural selection.

De ovi mammalium et hominis genesi (On the Genesis of the Ovum of Mammals and of Men) is an 1827 pamphlet by Karl Ernst von Baer about the anatomical observation and description of the egg (ovum) of mammals, like dogs and humans. The pamphlet detailed evidence for the existence of the ovum at the beginning of the developmental process in mammals. Prior to von Baer's publication, there was much debate about how organisms develop, as some claimed that organisms grow from a corpuscular element already preformed in the body (preformationism), and others said that organisms developed from a fluid material undergoing a process of progressive formation (epigenesis). Researchers at the time struggled to observe the early stages of development, and those such as von Baer had to observe the phenomenon through microscopes and then provide interpretations of the phenomena they observed.