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@ARTICLE{Bryant1984-pa,
title = "Intrinsic and extrinsic control of growth in developing organs",
author = "Bryant, P J and Simpson, P",
abstract = "The growth rate and final size of developing organs is controlled
by organ-intrinsic mechanisms as well as by hormones and growth
factors that originate outside the target organ. Recent work on
Drosophila imagined discs and other regenerating systems has led
to the conclusion that the intrinsic growth-control mechanism
that controls regenerative growth depends on position-specific
interactions between cells and their neighbors, and that these
interactions also control pattern formation. According to this
interpretation, local growth by cell proliferation is stimulated
when cells with disparate positional information are confronted
as a result of grafting or wound healing. This local growth leads
to intercalation of cells with intervening positional values
until the positional information discontinuity is eliminated.
When all discontinuities have been eliminated from a positional
field, growth stops. In this article we consider the possibility
that organ growth during normal development may be controlled by
an intercalation mechanism similar to that proposed for
regenerative growth. Studies of imaginal disc growth are
consistent with this suggestion, and in addition they show that
the cell interactions thought to control growth are independent
of cell lineage. Developing organs of vertebrates also show
intrinsic growth-control mechanisms, as demonstrated by the
execution of normal growth programs by immature organs that are
transplanted to fully grown hosts or to hosts with genetically
different growth parameters. Furthermore, these organ-intrinsic
mechanisms also appear to be based on position-specific cell
interactions, as suggested by the growth stimulation seen after
partial extirpation or rearrangement by grafting. In organs of
most adult vertebrates, the organ-intrinsic growth-control system
seems to be suppressed as shown by the loss of regenerative
ability, although it is clearly retained in the limbs, tails and
other organs of salamanders. The clearest example of an extrinsic
growth regulator is growth hormone, which plays a dominant role
along with insulin-like growth factors, thyroid hormone and sex
hormones in supporting the growth of bones and other organs in
postnatal mammals. These hormones do not appear to regulate
prenatal growth, but other hormones and insulin-like growth
factors may be important prenatally. The importance of other
growth factors in regulating organ growth in vivo remains to be
established. It is argued that both intrinsic and extrinsic
factors control organ growth, and that there may be important
interactions between the two types of control during development.",
journal = "Q Rev Biol",
volume = 59,
number = 4,
pages = "387--415",
month = dec,
year = 1984,
address = "United States",
language = "en"
}
@ARTICLE{Jun2018-wz,
title = "Fundamental principles in bacterial physiology-history, recent
progress, and the future with focus on cell size control: a
review",
author = "Jun, Suckjoon and Si, Fangwei and Pugatch, Rami and Scott,
Matthew",
abstract = "Bacterial physiology is a branch of biology that aims to
understand overarching principles of cellular reproduction. Many
important issues in bacterial physiology are inherently
quantitative, and major contributors to the field have often
brought together tools and ways of thinking from multiple
disciplines. This article presents a comprehensive overview of
major ideas and approaches developed since the early 20th century
for anyone who is interested in the fundamental problems in
bacterial physiology. This article is divided into two parts. In
the first part (sections 1-3), we review the first 'golden era'
of bacterial physiology from the 1940s to early 1970s and provide
a complete list of major references from that period. In the
second part (sections 4-7), we explain how the pioneering work
from the first golden era has influenced various rediscoveries of
general quantitative principles and significant further
development in modern bacterial physiology. Specifically, section
4 presents the history and current progress of the 'adder'
principle of cell size homeostasis. Section 5 discusses the
implications of coarse-graining the cellular protein composition,
and how the coarse-grained proteome 'sectors' re-balance under
different growth conditions. Section 6 focuses on physiological
invariants, and explains how they are the key to understanding
the coordination between growth and the cell cycle underlying
cell size control in steady-state growth. Section 7 overviews how
the temporal organization of all the internal processes enables
balanced growth. In the final section 8, we conclude by
discussing the remaining challenges for the future in the field.",
journal = "Rep Prog Phys",
volume = 81,
number = 5,
pages = "056601",
month = jan,
year = 2018,
language = "en"
}
@article{RHIND2021R1414,
title = {Cell-size control},
journal = {Current Biology},
volume = {31},
number = {21},
pages = {R1414-R1420},
year = {2021},
issn = {0960-9822},
doi = {https://doi.org/10.1016/j.cub.2021.09.017},
url = {https://www.sciencedirect.com/science/article/pii/S0960982221012574},
author = {Nicholas Rhind},
abstract = {Summary
A fundamental and still mysterious question in cell biology is “How do cells know how big they are?”. The fact that they do is evident from the strict maintenance of size homeostasis within populations of cells and has been verified by a variety of creative experiments over the past 100 years. An increasingly sophisticated understanding of cell-cycle-control mechanisms and innovations in cell imaging and analysis tools have allowed recent progress in proposing and testing models of cell-size control. Nonetheless, a biochemical understanding of how proposed cell-size mechanisms might work is only beginning to be developed. This primer introduces the field of cell-size control and discusses some of the questions that are yet to be answered.}
}
@ARTICLE{Shraiman2005-wl,
title = "Mechanical feedback as a possible regulator of tissue growth",
author = "Shraiman, Boris I",
abstract = "Regulation of cell growth and proliferation has a fundamental
role in animal and plant development and in the progression of
cancer. In the context of development, it is important to
understand the mechanisms that coordinate growth and patterning
of tissues. Imaginal discs, which are larval precursors of fly
limbs and organs, have provided much of what we currently know
about these processes. Here, we consider the mechanism that is
responsible for the observed uniformity of growth in wing
imaginal discs, which persists in the presence of gradients in
growth inducing morphogens in spite of the stochastic nature of
cell division. The phenomenon of ``cell competition,'' which
manifests in apoptosis of slower-growing cells in the vicinity of
faster growing tissue, suggests that uniform growth is not a
default state but a result of active regulation. How can a patch
of tissue compare its growth rate with that of its surroundings?
A possible way is furnished by mechanical interactions. To
demonstrate this mechanism, we formulate a mathematical model of
nonuniform growth in a layer of tissue and examine its mechanical
implications. We show that a clone growing faster or slower than
the surrounding tissue is subject to mechanical stress, and we
propose that dependence of the rate of cell division on local
stress could provide an ``integral-feedback'' mechanism
stabilizing uniform growth. The proposed mechanism of growth
control is not specific to imaginal disc growth and could be of
general relevance. Several experimental tests of the proposed
mechanism are suggested.",
journal = "Proc Natl Acad Sci U S A",
volume = 102,
number = 9,
pages = "3318--3323",
month = feb,
year = 2005,
language = "en"
}
@article{AEGERTERWILMSEN2007318,
title = {Model for the regulation of size in the wing imaginal disc of Drosophila},
journal = {Mechanisms of Development},
volume = {124},
number = {4},
pages = {318-326},
year = {2007},
issn = {0925-4773},
doi = {https://doi.org/10.1016/j.mod.2006.12.005},
url = {https://www.sciencedirect.com/science/article/pii/S0925477306002206},
author = {Tinri Aegerter-Wilmsen and Christof M. Aegerter and Ernst Hafen and Konrad Basler},
keywords = {Organ size, Growth control, Wing disc, , Dpp, Mechanical forces, Computer simulations},
abstract = {For animal development it is necessary that organs stop growing after they reach a certain size. However, it is still largely unknown how this termination of growth is regulated. The wing imaginal disc of Drosophila serves as a commonly used model system to study the regulation of growth. Paradoxically, it has been observed that growth occurs uniformly throughout the disc, even though Decapentaplegic (Dpp), a key inducer of growth, forms a gradient. Here, we present a model for the control of growth in the wing imaginal disc, which can account for the uniform occurrence and termination of growth. A central feature of the model is that net growth is not only regulated by growth factors, but by mechanical forces as well. According to the model, growth factors like Dpp induce growth in the center of the disc, which subsequently causes a tangential stretching of surrounding peripheral regions. Above a certain threshold, this stretching stimulates growth in these peripheral regions. Since the stretching is not completely compensated for by the induced growth, the peripheral regions will compress the center of the disc, leading to an inhibition of growth in the center. The larger the disc, the stronger this compression becomes and hence the stronger the inhibiting effect. Growth ceases when the growth factors can no longer overcome this inhibition. With numerical simulations we show that the model indeed yields uniform growth. Furthermore, the model can also account for other experimental data on growth in the wing disc.}
}
@ARTICLE{Hufnagel2007-nt,
title = "On the mechanism of wing size determination in fly development",
author = "Hufnagel, Lars and Teleman, Aurelio A and Rouault, Herv{\'e} and
Cohen, Stephen M and Shraiman, Boris I",
abstract = "A fundamental and unresolved problem in animal development is the
question of how a growing tissue knows when it has achieved its
correct final size. A widely held view suggests that this process
is controlled by morphogen gradients, which adapt to tissue size
and become flatter as tissue grows, leading eventually to growth
arrest. Here, we present evidence that the decapentaplegic (Dpp)
morphogen distribution in the developing Drosophila wing imaginal
disk does not adapt to disk size. We measure the distribution of
a functional Dpp-GFP transgene and the Dpp signal transduced by
phospho-Mad and show that the characteristic length scale of the
Dpp profile remains approximately constant during growth. This
finding suggests an alternative scenario of size determination,
where disk size is determined relative to the fixed morphogen
distribution by a certain threshold level of morphogen required
for growth. We propose that when disk boundary reaches the
threshold the arrest of cell proliferation throughout the disk is
induced by mechanical stress in the tissue. Mechanical stress is
expected to arise from the nonuniformity of morphogen
distribution that drives growth. This stress, through a negative
feedback on growth, can compensate for the nonuniformity of
morphogen, achieving uniform growth with the rate that vanishes
when disk boundary reaches the threshold. The mechanism is
demonstrated through computer simulations of a tissue growth
model that identifies the key assumptions and testable
predictions. This analysis provides an alternative hypothesis for
the size determination process. Novel experimental approaches
will be needed to test this model.",
journal = "Proc Natl Acad Sci U S A",
volume = 104,
number = 10,
pages = "3835--3840",
month = feb,
year = 2007,
language = "en"
}
@article{PhysRevLett.129.048102,
title = {Instabilities and Geometry of Growing Tissues},
author = {Grossman, Doron and Joanny, Jean-Francois},
journal = {Phys. Rev. Lett.},
volume = {129},
issue = {4},
pages = {048102},
numpages = {5},
year = {2022},
month = {Jul},
publisher = {American Physical Society},
doi = {10.1103/PhysRevLett.129.048102},
url = {https://link.aps.org/doi/10.1103/PhysRevLett.129.048102}
}
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thesis_notes.bbl 0 → 100644
\begin{thebibliography}{1}
\bibitem{AEGERTERWILMSEN2007318}
Tinri Aegerter-Wilmsen, Christof~M. Aegerter, Ernst Hafen, and Konrad Basler.
\newblock Model for the regulation of size in the wing imaginal disc of
drosophila.
\newblock {\em Mechanisms of Development}, 124(4):318--326, 2007.
\bibitem{Bryant1984-pa}
P~J Bryant and P~Simpson.
\newblock Intrinsic and extrinsic control of growth in developing organs.
\newblock {\em Q Rev Biol}, 59(4):387--415, December 1984.
\bibitem{PhysRevLett.129.048102}
Doron Grossman and Jean-Francois Joanny.
\newblock Instabilities and geometry of growing tissues.
\newblock {\em Phys. Rev. Lett.}, 129:048102, Jul 2022.
\bibitem{Hufnagel2007-nt}
Lars Hufnagel, Aurelio~A Teleman, Herv{\'e} Rouault, Stephen~M Cohen, and
Boris~I Shraiman.
\newblock On the mechanism of wing size determination in fly development.
\newblock {\em Proc Natl Acad Sci U S A}, 104(10):3835--3840, February 2007.
\bibitem{Jun2018-wz}
Suckjoon Jun, Fangwei Si, Rami Pugatch, and Matthew Scott.
\newblock Fundamental principles in bacterial physiology-history, recent
progress, and the future with focus on cell size control: a review.
\newblock {\em Rep Prog Phys}, 81(5):056601, January 2018.
\bibitem{RHIND2021R1414}
Nicholas Rhind.
\newblock Cell-size control.
\newblock {\em Current Biology}, 31(21):R1414--R1420, 2021.
\bibitem{Shraiman2005-wl}
Boris~I Shraiman.
\newblock Mechanical feedback as a possible regulator of tissue growth.
\newblock {\em Proc Natl Acad Sci U S A}, 102(9):3318--3323, February 2005.
\end{thebibliography}
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[1
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Chapter 1.
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Chapter 2.
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Chapter 3.
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Chapter 4.
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Chapter 5.
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Chapter 6.
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Appendix A.
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Appendix B.
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\contentsline {chapter}{\numberline {1}Size regulation in living systems}{2}%
\contentsline {chapter}{\numberline {2}Cell size regulation}{3}%
\contentsline {section}{\numberline {2.1}Evidence}{3}%
\contentsline {section}{\numberline {2.2}Cell size control mechanisms: Sizers, Adders and Timers}{3}%
\contentsline {section}{\numberline {2.3}Modeling sizers, adders and timers}{3}%
\contentsline {chapter}{\numberline {3}Tissue size regulation}{4}%
\contentsline {section}{\numberline {3.1}Existing models}{4}%
\contentsline {subsection}{\numberline {3.1.1}Signaling-based models}{4}%
\contentsline {subsection}{\numberline {3.1.2}Mechanical models}{4}%
\contentsline {chapter}{\numberline {4}A minimal mechanical model}{5}%
\contentsline {section}{\numberline {4.1}No signaling on fixed boundaries}{6}%
\contentsline {section}{\numberline {4.2}Moving boundaries}{6}%
\contentsline {section}{\numberline {4.3}Controlling growth}{6}%
\contentsline {chapter}{\numberline {5}Model on fixed boundaries}{7}%
\contentsline {chapter}{\numberline {6}Model on moving boundaries}{9}%
\contentsline {section}{\numberline {6.1}Diffusion on fixed boundaries becomes diffusion-advection on moving boundaries}{9}%
\contentsline {section}{\numberline {6.2}Conservation laws}{9}%
\contentsline {section}{\numberline {6.3}Exact solution for advection diffusion equation on a domain growing exponentially}{10}%
\contentsline {section}{\numberline {6.4}Exact solution for advection diffusion equation on a domain growing exponentially till $t=t_c$}{11}%
\contentsline {chapter}{\numberline {A}Visco-elastic models}{12}%
\contentsline {section}{\numberline {A.1}Linear visco-elasticity}{12}%
\contentsline {section}{\numberline {A.2}Models of linear viscoleasticity}{12}%
\contentsline {subsection}{\numberline {A.2.1}Maxwell model}{12}%
\contentsline {subsection}{\numberline {A.2.2}Kelvin-Voigt model}{12}%
\contentsline {chapter}{\numberline {B}Proof of the Reynolds Transport Theorem}{14}%
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