Sean C. Sleight and Herbert M. Sauro: Coli, Propagated With All Colors Turned On. Our Results Using

DOI: http://dx.doi.org/10.
7551/978-0-262-31050-5-ch063
Design and construction of a prototype CMY (Cyan-Magenta-Yellow) genetic circuit

as a mutational readout device to measure evolutionary stability dynamics and
determine design principles for robust synthetic systems
Sean C. Sleight1 and Herbert M. Sauro1
1 University of Washington, Dept. of Bioengineering, Seattle, WA 98195
sleight@uw.edu
Abstract (Cooper et al. 2001; Colosimo et al. 2005; Sleight et al. 2008;
Downloaded from http://direct.mit.edu/isal/proceedings-pdf/alife2012/24/481/1901038/978-0-262-31050-5-ch063.pdf by guest on 14 January 2022

Synthetic biology is the engineering discipline for constructing Meyer et al. 2010; Toprak et al. 2012).
novel organisms with functions that do not exist in nature. We For a number of reasons, synthetic biology offers a
recently engineered a prototype CMY genetic circuit that powerful system for studying evolution in the laboratory.
visually produces cyan, magenta, and yellow colors Synthetic biologists assemble genetic circuits and metabolic
independently and in combination using different inducer pathways from individual genetic “parts” (Knight, 2003;
molecules. Since the production of each color can be Shetty et al., 2008; Sleight et al., 2010a), normally encoded on
independently controlled, this allows for the production of a plasmids (Elowitz & Leibler, 2000; Gardner et al., 2000; Basu
spectrum of colors that can be visualized in normal light et al., 2005; Levskaya et al., 2005; Entus et al., 2007;
conditions, and each color can be quantified using fluorescence Anderson et al., 2007; Sleight et al., 2010b), but also on the
measurements. We performed an evolution experiment to
chromosome (Tyo et al., 2009; Gibson et al., 2010). Each
measure the evolutionary stability dynamics of this prototype
CMY genetic circuit in 88 replicate populations of Escherichia genetic circuit has a unique metabolic load associated with it,
coli, propagated with all colors turned on. Our results using due to the production of foreign proteins, and as a result has a
particular inducer concentrations show that all 88 replicate unique fitness. Any cell in the population that removes this
populations change from a dark, green-brown color to a cyan- metabolic load through a loss-of-function mutation normally
ish color after only 40 generations. In order to visualize the has a large fitness increase (unpublished results). Because of
results of this experiment, we washed and concentrated the cells this large fitness differential between functional and
from each population into a different well of a 384-well plate at nonfunctional cells, evolution occurs rapidly due to the
different evolutionary timepoints. The color change seen functional cells in the population being outcompeted by
visually is confirmed with quantitative data that demonstrates
nonfunctional cells. There are several examples of genetic
the loss-of-function of both magenta and yellow colors with
variation between replicate populations. We sequenced a single circuits (You et al., 2004; Balagaddé et al., 2005; Canton et
clone from four independently evolved populations and all al., 2008; Sleight et al., 2010b) and metabolic pathways (Yoon
clones have the same loss-of-function deletion mutation et al., 2007; Philip et al., 2009; Tyo et al., 2009) that have lost
between homologous transcriptional terminators that removes function over evolutionary time. The loss-of-function
the magenta and yellow expression cassettes. This parallel mutations can easily be determined by comparing the original
evolution was somewhat expected from results of previous plasmid sequence with the plasmid extracted from individual
work, but we expect that randomized and re-engineered evolved clones. One disadvantage to studying evolutionary
versions of this circuit without repeats will produce more dynamics in plasmids is that there may be unknown mutations
divergent results due to more stochastic loss-of-function that occur on the host chromosome, but transforming the
mutations. This prototype CMY circuit serves as a mutational
readout device and allows for a colorimetric and quantitative original and mutant plasmids back into plasmid-less host,
demonstration of evolution in action using synthetic biology. allows for determination of the mutant plasmid phenotype
(Sleight et al., 2010b). Thus, synthetic biologists are able to
engineer different genetic circuits with control over the exact
Introduction DNA sequence, transform them into a host organism of
choice, perform evolution experiments that often occur over
The field of experimental evolution uses controlled short timescales, and determine the exact mutations
experiments for studying evolutionary dynamics over short or responsible for evolutionary adaptation.
long timescales in the laboratory (Elena and Lenski 2003). As genetic circuits get more complex, it becomes
Evolution experiments in the laboratory normally involve increasingly important to understand the evolutionary stability
propagating bacteria or other microbes over multiple dynamics of large circuits with high metabolic loads. With this
generations in certain environmental conditions to understand goal in mind, we recently engineered a prototype CMY
the phenotypic and genetic differences between evolved (Cyan-Magenta-Yellow) genetic circuit (Figure 1) to study
strains and their progenitors (Riehle, Bennett, and Long 2005; the evolutionary stability dynamics of a three-gene circuit.
Herring et al. 2006; Schoustra et al. 2006; Sleight and Lenski After various iterations, we found that this circuit expresses
2007). These experiments also allow for the study of parallel colors best using medium-copy plasmids (instead of high-
or divergent evolutionary processes at the genetic and copy which cause instability) and strong ribosomal binding
phenotypic levels between replicate evolved populations. sites. Each color in the circuit can be turned on independently
Parallel evolution occurs when multiple evolved populations and in combination using different inducer molecules,
derived from the same ancestor converge on a similar producing a spectrum of different colors. The color can be
phenotype and is a strong indicator of evolutionary adaptation
© 2012 Massachusetts Institute of Technology Artificial Life 13: 481–488

Design and construction of a prototype CMY (Cyan-Magenta-Yellow) genetic circuit as a mutational readout device to
measure evolutionary stability dynamics and determine design principles for robust synthetic systems
seen visually in normal light conditions when the cells are colony was grown for 24 hours in 5 mL LB + 50 µg/mL
pelleted, washed, and resuspended in water. Importantly, each kanamycin in a test tube grown at 37°C shaking at 250 RPM.
color can also be quantified by measuring fluorescence. These Eight populations were inoculated (1:1000 dilution) into 5
colorimetric and quantitative methods allow for a mutational mL LB + 50 µg/mL kanamycin in test tubes grown at 37°C
“readout” of circuit function at any evolutionary timepoint. In shaking at 250 RPM and supplemented with different inducers
this study, we independently evolved 88 replicate populations and molecules. Expression of LacZ was induced with 0.02%
to understand the mutational robustness of this prototype arabinose from the pBAD (I0500) promoter. X-gal was added
circuit and whether the replicate populations would evolve in to the media to visualize the presence of beta-galactosidase,
parallel or divergently. These results will allow for a better expressed from the lacZ coding sequence. X-gal is cleaved by
understanding of how to engineer robust synthetic systems. β-galactosidase yielding galactose and 5-bromo-4-chloro-3-
hydroxyindole. The latter is then oxidized into 5,5'-
dibromo-4,4'-dichloro-indigo, an insoluble blue product. The
Materials and Methods molecule 4-Methylumbelliferyl beta-D-galactopyranoside
(MUG) dissolved in DMSO was added to the media to
Circuit engineering and use of strains. The prototype quantify the concentration of LacZ using fluorescence

CMY circuit (Figure 1) was engineered from DNA obtained measurements (Vidal-Aroca et al. 2006). Green Fluorescent
from the Registry of Standard Biological Parts Protein (GFP) was induced with Anhydrotetracycline (aTc)
(partsregistry.org) using the Clontech In-Fusion PCR Cloning from the pTetR (R0040) promoter. Monomeric Red
Kit, with the specific methods described previously (Sleight et Fluorescent Protein (mRFP) was induced with Isopropyl-beta-
al., 2010a). This circuit was cloned in the pSB3K3 plasmid, a D-thiogalactopyranoside (IPTG) from the pLacI (R0010)
medium copy number plasmid (20-30 plasmids/cell) with a promoter. The following inducers and molecules were added
to eight controls according to the table below (+ indicates the
addition of the inducers and molecules listed in the column
and – indicates absence of the same inducers and molecules).
Results of this experiment are shown in Figure 2.
Inducers and Arabinose (0.02%), aTc IPTG

molecules X-gal (10 µg/mL) (1 X 10-4 M)
(20 µg/ml), MUG
Figure 1. Design and regulation of the prototype CMY genetic
(2 µg/mL)
circuit. The CMY circuit expresses three proteins (LacZ, GFP, and
Control #1 + - -
mRFP) independently and in combination using different inducer
molecules. The inducer molecule arabinose (Ara) binds to the AraC Control #2 - + -
protein (encoded on the plasmid) and activates expression of LacZ Control #3 - - +
from the pBAD promoter. The addition of X-gal in the media allows
for the visualization of beta-galactosidase (LacZ) expression since Control #4 + + -
LacZ cleaves this molecule to produce a blue color (seen as cyan in Control #5 - + +
normal light conditions if a particular concentration is used). The
Control #6 + - +
addition of MUG in the media allows for the quantification of LacZ.
The cells turn yellow in normal light conditions when the inducer aTc Control #7 + + +
binds to TetR (encoded on the chromosome) and derepresses Control #8 - - -
expression of GFP from the pTetR promoter. To produce a visual red
color, IPTG binds to LacI (encoded on the chromosome) and
derepresses expression of mRFP from the pLacI promoter. The Evolution experiment. The Z1 strain transformed with the
genetic symbols (taken from SBOL visual) represent promoters CMY plasmid was streaked out from a freezer stock of a
(broken arrows with activation or repression), ribosome binding sites culture frozen with 15% glycerol and stored at -80°C. One
(half ovals), coding sequences (forward arrows), and transcriptional colony was grown for 24 hours in 5 mL LB + 50 µg/mL
terminators (“T” symbol). Lines ending with a filled circle indicate kanamycin in a test tube grown at 37°C shaking at 250 RPM.
activation and lines ending with a perpendicular line indicate Eight-eight identical populations were inoculated from this
repression. See details in the Materials and Methods section. culture (1:1000 dilution) into 1.5 mL LB + 50 µg/mL
p15A pMR101-derived replication origin and kanamycin kanamycin and supplemented with 0.02% arabinose, 20 µg/ml
resistance gene (Lutz & Bujard, 1997). This plasmid was X-gal, 2 µg/mL MUG, 10 µg/mL aTc, and 1 X 10-4 M IPTG in
transformed into MG1655 Z1 (Sleight et al., 2010b) which an Eppendorf deep 96-well plate sealed with a Thermo
constitutively overexpresses LacI and TetR from the Scientific gas permeable membrane for maximum oxygen
chromosome. The AraC protein is expressed from a diffusion. These cultures were grown at 37°C shaking at 250
constitutive promoter on I0500 in the reverse direction, RPM and propagated every 24 hours achieve about 10
whereas LacZ is expressed from the pBAD promoter on I0500 generations per day (log2 1000 = 9.97).
in the forward direction.
Cell density and fluorescence measurements. Every
Independent and combinatorial expression of each 24 hours, cell density (OD600) and fluorescence of evolved
color on the CMY circuit. The Z1 strain transformed with populations were measured in a Tecan Infinite M200 Pro
the CMY plasmid was streaked out from a freezer stock of a fluorescence plate reader. The measurement timepoint chosen
culture frozen with 15% glycerol and stored at -80°C. One was every 24 hours because the rate of change of fluorescent
protein expression is close to steady-state. Evolved
482 Artificial Life 13

populations thus spend about 8-12 hours in lag or exponential A .
phase and the remaining time in stationary phase. For each
timepoint, all populations were thoroughly mixed and 200 µl
was transferred into a black, clear-bottom 96-well plate
(Costar). Fluorescence was measured for LacZ expression
using 360nm excitation/460nm emission wavelengths, GFP
using 485 nm excitation/516 nm emission wavelengths, and
mRFP using 584 nm excitation/620 nm emission wavelengths.
Fluorescence for each color was then divided by OD600 to
measure the normalized expression (Fluorescence/OD600).
Plasmid sequencing. After 40 generations, four evolved

populations were streaked out on LB + 50 µg/mL kanamycin
agar plates. One clone from each population was grown in 5
mL LB + 50 µg/mL kanamycin for 24 hours at 37°C shaking

at 250 RPM. Plasmids were extracted using the Qiagen
Miniprep Kit and submitted to the Genewiz sequencing
facility for sequencing. Purified plasmid DNA was sequenced
using VF2/VR primers specific to the pSB3K3 vector (about B .
100 bp on either side of the circuit) and internal primers
specific to the circuit.
Visualizing cell color in controls and evolved

populations. The cell color for controls #1-8 were visualized
by centrifuging 5 mL test tubes in a Sorvall Legend 23R
centrifuge at 3000 RPM for 10 minutes at 4°C, removing the
supernatant, washing with 500 µL of water, centrifuging again
at 3000 RPM for 10 minutes at 4°C, removing the
supernatant, then resuspending the cells in 100 µL of water.
In Figure 3a, 50 µL of resuspended cells were added to
individual wells in a clear 384-well plate. The cells were
incubated in the plate for 24 hours to allow the cells to
“develop” color on the bottom of the well. The plate was
photographed upside down in normal light conditions. In
Figure 3b, 5 µL of resuspended cells were added to individual
wells in a 1536-well plate and visualized upside down using a C.
UV transilluminator without a filter. In Figure 6, Controls
#1-8 were added at each evolutionary timepoint as a color
reference for circuit function, but a lower concentration of
cells (1.5 mL cells / 300 µL of water) was used compared to
the image shown in Figure 3a, resulting in a lighter color.
For evolved populations, every 10 generations the deep 96-
well plate was centrifuged at 2000 RPM for 10 minutes at
4°C, removing the supernatant, washing with 500 µL of water,
centrifuging again at 2000 RPM for 10 minutes at 4°C,
removing the supernatant, then resuspending the cells in 100
µL of water. 50 µL of the resuspended cells were visualized
as described above for Figure 3a.
Control 1 2 3 4 5 6 7 8
Figure 2. Independent and combinatorial expression of each color

from the CMY circuit. Eight control cultures grown with different
inducers and molecules (see Materials and Methods) were measured
for (A) expression of LacZ, (B) expression of GFP, and (C)
expression of mRFP. The inducers Ara, aTc, and IPTG express LacZ,
GFP, and mRFP, respectively. Error bars represent one standard
deviation from the mean of three independent replicates.

A. A.
7 6 5 4 3 2 1 8
B.

B.
Figure 3. Visualization of CMY circuit colors under (A) normal light

and (B) UV light. In normal light conditions, the color for controls
#1-8 are visualized as described in Materials and Methods and the
control numbers are labeled underneath each colored square well. In
UV light, the color for controls #1-6 are visualized as described in
Materials and Methods, as follows: “A” is written with control #3,
“Life” with control #1, “13” with control #2, “evolution” with control
#5, “in” with control #6, and “action” with control #4.
Results
Independent and combinatorial expression of each C.
color from the CMY circuit. To test the functionality of
the CMY circuit, we first performed a control experiment to
measure expression of each color using different inducers and
molecules (see Materials and Methods for details). The
results of this experiment is shown in Figure 2. Starting with
Figure 2a, the results show that LacZ is expressed about 10-
fold above background levels with addition of arabinose, but
not with aTc or IPTG. With other inducer combinations, LacZ
is only expressed with arabinose, but to varying levels. This
indicates that expression of other genes in the circuit affect
expression of LacZ, possibly due to competition for
expression and metabolic load. In Figure 2b, GFP is only
expressed with the addition of aTc and is about 100-fold
above background levels with other inducers. Like LacZ,
GFP expression is also affected by expression of other genes
in the circuit. Figure 2c shows that mRFP is expressed only
with the addition of IPTG and is over 10-fold above
background levels with other inducers. Overall, the results
indicate the independent and combinatorial expression of each
color in the CMY circuit, but expression levels are affected by Figure 4. Evolutionary stability dynamics of the CMY circuit.
different combinations of inducers. Figure 3 shows visually Normalized fluorescence on the y-axis is plotted against the number
that distinct colors are produced with each combination of of generations for (A) MUG fluorescence (LacZ), (B) GFP
inducer used, demonstrating combinatorial expression. Note fluorescence, and (C) mRFP fluorescence. Error bars represent the
that GFP appears yellow visually in Figure 3a, but is green in standard deviation from the mean of 88 independent replicates.
Figure 3b under UV light.

Evolutionary stability dynamics of the CMY circuit.
Next, we performed an evolution experiment to measure the
evolutionary stability dynamics of the CMY circuit and
determine whether replicate populations evolved in parallel or
divergently. For this experiment, we evolved 88 replicate
populations in conditions where all colors in the circuit are
turned on to increase the metabolic load and thereby
maximize evolutionary processes. The results of this
experiment are shown in Figure 4. Figure 4a shows the
evolutionary stability dynamics of LacZ which remains
relatively constant for 40 generations, with slightly increased Figure 5. Dominant loss-of-function mutation in the CMY circuit.
expression at generation 40. In contrast, Figures 4b and 4c Individual clones from four independently evolved populations all
show that both GFP and mRFP expression is constant for have the same loss-of-function mutation: a deletion between repeated
B0015 terminators that effectively removes the GFP and mRFP
about 10 generations with fluctuations, then both dramatically
expression cassettes. The mutant clones can still express LacZ.
decrease by generation 20. Both GFP and mRFP expression

continues to decrease slowly, but have not dropped to
background levels by generation 40.
Loss-of-function mutation in the CMY circuit. To

determine the mutation responsible for decreased GFP and Discussion and Future Directions
mRFP expression, four evolved populations from the 40 In this study, we demonstrate the design and engineering of a
generation timepoint were streaked out and a single clone functional CMY circuit, where each color in the circuit can be
from each was grown overnight for plasmid extraction. The expressed independently and in combination. To measure the
mutant plasmid was shown to have a deletion between evolutionary stability dynamics of this prototype circuit, we
repeated B0015 terminators (129 bp) that effectively removed evolved 88 independent populations with all colors turned on.
both the GFP and mRFP expression cassettes (Figure 5). The We observed striking parallel evolution in the color change of
sequencing data was very clean, indicating that the plasmids the evolved populations that is in agreement with the
within each clone sequenced were likely identical or nearly quantitative measurements. The dominant mutation that we
identical (e.g. noisy sequencing data may have indicated a found in four of the populations is a deletion between repeated
mixture of different plasmids). This mutation was terminators that effectively removes the GFP and mRFP
unsurprising since similar deletions between repeated expression cassettes (note that there is no significant
terminators have occurred in other circuits we have studied homology between GFP and mRFP). Although only clones in
(Sleight et al., 2010b). Although this result was somewhat four of the 88 populations were sequenced, it is likely that the
expected, we still did not know exactly how a three-gene other populations had the same loss-of-mutation. Incidentally,
circuit would lose function. Interestingly, in a pilot evolution a previous study found that a circuit with repeated B0015
study, an earlier version of this circuit with different genetic terminators had a deletion between these sequences even
parts that was transformed into a different strain, using when using a strain with a recA mutation and therefore
replication slippage alone can cause this common mutation
different inducer concentrations than reported here, lost the
(Canton et al., 2008). Our previous work on the evolutionary
cyan function first.
stability of another genetic circuit with repeated B0015
terminators showed that nine out of nine populations lost
Visualizing color variation between 88 replicate function due to the same deletion between these repeated
evolved populations. We expected to see color differences sequences, but re-engineering the circuit to have non-
between replicate populations over evolutionary time. In homologous terminators increases its evolutionary stability
order to visualize the color of each evolved population at (Sleight et al., 2010b). The prototype CMY circuit in this
different evolutionary timepoints, we washed and resuspended study had repeated B0015 terminators only because this was
the cells in water, then added these cells to individual wells in the first version of the circuit tested to determine if the circuit
a 384-well plate. The results are shown in Figure 6. While was functional. We expect some future versions of this circuit
there will be variation in color due to experimental methods to evolve more divergently (produce a wide variety of colors)
(e.g. different number of cells put in each well) as well as due to the absence of repeated sequences. Also, it may be
variation due to the stochastic nature of evolutionary interesting to understand how the absence of different
dynamics, the photos of each plate at different timepoints inducers in the media changes the evolutionary stability
clearly show a color change from a dark greenish-brown to dynamics and loss-of-function mutations in future versions of
cyan color over the course of 40 generations. The sharpest the circuit. We have recently developed an assembly method
transition in color is seen between generations 10 and 20, to randomize parts (e.g. promoters, coding sequences,
matching the quantitative data (Figure 4) closely. Generation transcriptional terminators) to generate different combinations
20 cells are a dark cyan color since there is still some GFP and of genetic circuits. We will use this method to shuffle parts to
mRFP being expressed. By generation 40, the cells appear to engineer various CMY circuits, then use a directed evolution
approach to evolve selected circuits individually and pooled to
match the cyan color of the circuit when only LacZ is
determine which versions of the circuit are most
expressed. The cyan color would likely slowly fade away if
evolutionarily robust. We expect that some circuit variants
the evolution experiment was continued beyond 40 will evolve in parallel and some will evolve divergently due to
generations. genotypic (e.g. repeat sequences, GC content, specific genetic
elements) and phenotypic (e.g. metabolic load due to the

expression of foreign proteins individually and in A. D .

B. E.
C . Figure 6. Color variation in evolved CMY circuit populations. Cells

from each of the 88 independently evolved populations at different
evolutionary timepoints were washed and resuspended in water, then
added to individual wells in a 384-well plate to spell Alife13. Each
plate represents (A) Generation 0, (B) Generation 10, (C) Generation
20, (D) Generation 30, and (E) Generation 40. The bottom right
corner shows eight colored wells that represent the controls #1-8 (see
Materials and Methods). The #8 control (unpigmented cells with no
inducers) is shown on the left, then the #1-7 controls are shown going
from left to right. Since the same cells were used for all controls on
all plates, the color fades in the later timepoints.
combination) differences. The evolutionary stability dynamics

and loss-of-function mutations will be measured in order to
better understand parallel/divergent evolutionary processes
and determine design principles for robust synthetic systems.
Ideally a selective pressure will be used to maintain function
of circuit components, but when that is not possible, the next
best method is to lower expression level and mutation rate to a
level that maintains function for as long as possible. If a high
expression level is needed, then the next best method is to
rationally design circuits that mutate in a predictable manner
such that multiple versions of the circuit can turn on, then off
via mutation. With this goal in mind, we also aim to identify a
CMY circuit variant that will act as an “evolutionary timer
circuit” for industrial applications when timed functions are
needed, such that one color loses function after x generations,
then a second color is lost after another y generations, and
finally a third color loses function after z generations.

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ananatis.” Applied microbiology and biotechnology 74 (1)
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killing.” Nature 428 (6985): 868-871.

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Real-Time Redesign Section 3: Start Small Step 1: Design and Run a Pilot
Activity: Visualize and Build a Prototype

Activity: Visualize Framing
and Build a Now that you’ve selected a solution to solve your
Prototype
problem of practice, it’s time to build out the details by
creating a prototype, or testable model of your solution.
The prototype will allow you to get feedback on the
Create a small-scale version of your idea and pilot it with a small group of stakeholders (e.g.,
solution students, teachers, parents, and/or classified staff). For
instance, if your solution was a new system for student-
led conferences, your prototype could be an agenda for
what those conferences might look like.
It can be tempting to build out a complete, complex

prototype, but it is important to keep it simple. A simpler
prototype is more conducive to getting quick stakeholder
feedback and to actually testing the concept in a “bite-
sized” way, allowing you to learn what is and is not
effective in the prototype to iterate as needed.
Ensuring Equity & Resiliency
Equity looks like building your prototype with a focus Resiliency looks like building a prototype that is clear and
on how your model will better meet the needs of each simple enough to be replicated and changed - especially
student in your system, especially those included based on quick stakeholder feedback. You may even look
in the empathy interview process. Equity also looks for ways to explicitly build flexibility and adaptability into
like sharing your prototype with students, families, your prototype (for example, through student choice).
teachers, and classified staff for feedback – and then
implementing changes based on that feedback. To
maximize stakeholder input, co-design the prototype with
the students, families, teachers, and classified staff you
interview.
This work is licensed under a Creative Commons Attribution 4.0 International License 55

Steps to Implementation
Suggested time: 2 hours
1. G
ather your team in a room (in-person or virtual) with access to a chalkboard or a digital alternative, such as Google
Jamboard; make sure your team has selected a solution to your problem of practice.
2. Given your solution, decide as a group what your prototype could look like or include.
• F
or example, if your solution is about standards-based grading, you may choose to prototype a new grading
rubric.
• F
or example, if your solution is about a new classroom schedule for more individualized student support, you may
choose to prototype a new lesson plan.
3. N
ext, brainstorm a definition of success for your prototype. Ask, “What do we believe will lead to our prototype
design and content being most successful?” Document ideas on a chalkboard or Jamboard.
• F
or example, if you are designing a new grading rubric, you might define success for your prototype as being
standards-aligned, using student-friendly language, and allowing for flexibility for teachers.
4. T
hen, build the prototype. Your team may work as a full group or you can break into small groups to design several
prototypes. Use the templates below for building a prototype; choose the template that aligns best with the format
of what you are trying to create (e.g., a visual of a new grading rubric is likely obvious, but a storyboard might also
help demonstrate how students experience using it).
• Storyboard
• Rapid Prototyping
• Get Visual
• Roleplay
5. If you break into small groups to create several prototypes, you will need to decide between the options (or take the
best aspects of each option to create a final version). One way to do this is to:
• Have each individual read each prototype and add a “+1” to indicate aspects that they like.
• Take time for group discussion; make real-time updates to the prototype(s) as you discuss:

Discussion Question Notes
What parts of each prototype did we like?
Was there one prototype that was a clear winner?

How could we combine the best parts of each
prototype into a new version?
How well does the prototype align to our definition of

success?
Where can we make final improvements or updates?
6. If you created one prototype as a group, take time for group discussion; make real-time updates as you discuss:
Discussion Question Notes
How well does the prototype align to our definition of

success?
How well does the prototype align with our original

problem statement?
Where can we make final improvements or updates?
7. Together, finalize your prototype.
• C
onsider getting interim feedback on this prototype by picking a handful of people to share the prototype with,
and prioritize getting feedback from the stakeholders you included earlier in this process. As a team, commit to
how you will collect and incorporate additional feedback.
8. Next, you will plan and execute a pilot test to put your prototype in action.
This activity was inspired by the Create a Prototype step of IDEO’s Design Thinking for Educators. The specific protocols
for building a prototype (Storyboard, Rapid Prototyping, Get Visual, Role Play) are IDEO’s method cards.
Prototype
A prototype is an early sample, model, or release of a product built
to test a concept or process.[1] It is a term used in a variety of
contexts, including semantics, design, electronics, and software
programming. A prototype is generally used to evaluate a new
design to enhance precision by system analysts and users.[2]
Prototyping serves to provide specifications for a real, working
system rather than a theoretical one.[3] In some design workflow
models, creating a prototype (a process sometimes called
materialization) is the step between the formalization and the
evaluation of an idea.[4]
A prototype can also mean a typical example of something such as

in the use of the derivation 'prototypical'.[5] This is a useful term in
identifying objects, behaviours and concepts which are considered
the accepted norm and is analogous with terms such as stereotypes
and archetypes.
Prototype signage on the Boise
The word prototype derives from the Greek πρωτότυπον Greenbelt, testing for rust, paint-
prototypon, "primitive form", neutral of πρωτότυπος prototypos, fastness, durability, etc.
"original, primitive", from πρῶτος protos, "first" and τύπος typos,
"impression".[1][6]
Contents
Types
Differences in creating a prototype vs. a final product
Characteristics and limitations of prototypes
Engineering sciences
Mechanical and electrical engineering
Electronics prototyping
Computer programming and computer science
Other programming/computing concepts
Data prototyping
Prototyping for Human-Computer Interaction A sign explaining prototype signage
Scale modeling
Architecture
Metrology
Natural sciences
See also
References
Types
Prototypes explore different aspects of an intended design:[7]
A proof-of-principle prototype serves to verify some key functional aspects of the intended
design, but usually does not have all the functionality of the final product.[8]
A working prototype represents all or nearly all of the functionality of the final product.[9]
A visual prototype represents the size and appearance, but not the functionality, of the
intended design. A form study prototype is a preliminary type of visual prototype in which
the geometric features of a design are emphasized, with less concern for color, texture, or
other aspects of the final appearance.[10]
A user experience prototype represents enough of the appearance and function of the
product that it can be used for user research.[11]
A functional prototype captures both function and appearance of the intended design,
though it may be created with different techniques and even different scale from final
design.[12][13]
A paper prototype is a printed or hand-drawn representation of the user interface of a
software product. Such prototypes are commonly used for early testing of a software design,
and can be part of a software walkthrough to confirm design decisions before more costly
levels of design effort are expended.[14]
Differences in creating a prototype vs. a final product

In general, the creation of prototypes will differ from creation of the final product in some fundamental
ways:
Material: The materials that will be used in a final product may be expensive or difficult to
fabricate, so prototypes may be made from different materials than the final product. In some
cases, the final production materials may still be undergoing development themselves and
not yet available for use in a prototype.
Process: Mass-production processes are often unsuitable for making a small number of
parts, so prototypes may be made using different fabrication processes than the final
product. For example, a final product that will be made by plastic injection molding will
require expensive custom tooling, so a prototype for this product may be fabricated by
machining or stereolithography instead. Differences in fabrication process may lead to
differences in the appearance of the prototype as compared to the final product.
Verification: The final product may be subject to a number of quality assurance tests to
verify conformance with drawings or specifications. These tests may involve custom
inspection fixtures, statistical sampling methods, and other techniques appropriate for
ongoing production of a large quantity of the final product. Prototypes are generally made
with much closer individual inspection and the assumption that some adjustment or rework
will be part of the fabrication process. Prototypes may also be exempted from some
requirements that will apply to the final product.
Engineers and prototype specialists attempt to minimize the impact of these differences on the intended role
for the prototype. For example, if a visual prototype is not able to use the same materials as the final
product, they will attempt to substitute materials with properties that closely simulate the intended final
materials.
Characteristics and limitations of prototypes

Engineers and prototyping specialists seek to understand the
limitations of prototypes to exactly simulate the characteristics of
their intended design.
It is important to realize that by their very definition, prototypes will

represent some compromise from the final production design. Due
to differences in materials, processes and design fidelity, it is
possible that a prototype may fail to perform acceptably whereas
the production design may have been sound. A counter-intuitive
idea is that prototypes may actually perform acceptably whereas the
A prototype of the Polish economy
production design may be flawed since prototyping materials and
hatchback car Beskid 106 designed
processes may occasionally outperform their production
in the 1980s
counterparts.
In general, it can be expected that individual prototype costs will be

substantially greater than the final production costs due to inefficiencies in materials and processes.
Prototypes are also used to revise the design for the purposes of reducing costs through optimization and
refinement.[15]
It is possible to use prototype testing to reduce the risk that a design may not perform as intended, however
prototypes generally cannot eliminate all risk. There are pragmatic and practical limitations to the ability of
a prototype to match the intended final performance of the product and some allowances and engineering
judgement are often required before moving forward with a production design.
Building the full design is often expensive and can be time-consuming, especially when repeated several
times—building the full design, figuring out what the problems are and how to solve them, then building
another full design. As an alternative, rapid prototyping or rapid application development techniques are
used for the initial prototypes, which implement part, but not all, of the complete design. This allows
designers and manufacturers to rapidly and inexpensively test the parts of the design that are most likely to
have problems, solve those problems, and then build the full design.
This counter-intuitive idea—that the quickest way to build something is, first to build something else—is
shared by scaffolding and Thomson's telescope rule.
Engineering sciences
In technology research, a technology demonstrator is a prototype serving as proof-of-concept and
demonstration model for a new technology or future product, proving its viability and illustrating
conceivable applications.
In large development projects, a testbed is a platform and prototype development environment for rigorous
experimentation and testing of new technologies, components, scientific theories and computational
tools.[16]
With recent advances in computer modeling it is becoming practical to eliminate the creation of a physical
prototype (except possibly at greatly reduced scales for promotional purposes), instead modeling all aspects
of the final product as a computer model. An example of such a development can be seen in Boeing 787
Dreamliner, in which the first full sized physical realization is made on the series production line. Computer
modeling is now being extensively used in automotive design, both for form (in the styling and
aerodynamics of the vehicle) and in function—especially for improving vehicle crashworthiness and in
weight reduction to improve mileage.
Mechanical and electrical engineering
The most common use of the word prototype is a functional, although experimental, version of a non-
military machine (e.g., automobiles, domestic appliances, consumer electronics) whose designers would
like to have built by mass production means, as opposed to a mockup, which is an inert representation of a
machine's appearance, often made of some non-durable substance.
An electronics designer often builds the first prototype from breadboard or stripboard or perfboard,
typically using "DIP" packages.
However, more and more often the first functional prototype is built on a "prototype PCB" almost identical
to the production PCB, as PCB manufacturing prices fall and as many components are not available in DIP
packages, but only available in SMT packages optimized for placing on a PCB.
Builders of military machines and aviation prefer the terms "experimental" and "service test".[17]
Electronics prototyping
In electronics, prototyping means building an actual circuit to a

theoretical design to verify that it works, and to provide a physical
platform for debugging it if it does not. The prototype is often
constructed using techniques such as wire wrapping or using
veroboard or breadboard, with the result being a circuit that is
electrically identical to the design but not physically identical to the
final product.[18]
Open-source tools like Fritzing exist to document electronic

A simple electronic circuit prototype
prototypes (especially the breadboard-based ones) and move
on a breadboard
toward physical production. Prototyping platforms such as Arduino
also simplify the task of programming and interacting with a
microcontroller.[19] The developer can choose to deploy their
invention as-is using the prototyping platform, or replace it with
only the microcontroller chip and the circuitry that is relevant to
their product.
A technician can quickly build a prototype (and make additions and

modifications) using these techniques, but for volume production it
is much faster and usually cheaper to mass-produce custom printed
circuit boards than to produce these other kinds of prototype
boards. The proliferation of quick-turn PCB fabrication and
assembly companies has enabled the concepts of rapid prototyping
to be applied to electronic circuit design. It is now possible, even
with the smallest passive components and largest fine-pitch Example of prototype in
packages, to have boards fabricated, assembled, and even tested in optoelectronics (Texas Instruments,
a matter of days. DLP Cinema Prototype System)
Computer programming and computer

science
Prototype software is often referred to as alpha grade, meaning it is the first version to run. Often only a few
functions are implemented, the primary focus of the alpha is to have a functional base code on to which
features may be added. Once alpha grade software has most of the required features integrated into it, it
becomes beta software for testing of the entire software and to adjust the program to respond correctly
during situations unforeseen during development.[20]
Often the end users may not be able to provide a complete set of application objectives, detailed input,
processing, or output requirements in the initial stage. After the user evaluation, another prototype will be
built based on feedback from users, and again the cycle returns to customer evaluation. The cycle starts by
listening to the user, followed by building or revising a mock-up, and letting the user test the mock-up, then
back. There is now a new generation of tools called Application Simulation Software which help quickly
simulate application before their development.[21]
Extreme programming uses iterative design to gradually add one feature at a time to the initial prototype.[22]
Other programming/computing concepts
In many programming languages, a function prototype is the declaration of a subroutine or function (and
should not be confused with software prototyping). This term is rather C/C++-specific; other terms for this
notion are signature, type and interface. In prototype-based programming (a form of object-oriented
programming), new objects are produced by cloning existing objects, which are called prototypes.[23]
The term may also refer to the Prototype Javascript Framework.
Additionally, the term may refer to the prototype design pattern.
Continuous learning approaches within organizations or businesses may also use the concept of business or
process prototypes through software models.
The concept of prototypicality is used to describe how much a website deviates from the expected norm,
and leads to a lowering of user preference for that site's design.[24]
Data prototyping
A data prototype is a form of functional or working prototype.[25] The justification for its creation is usually
a data migration, data integration or application implementation project and the raw materials used as input
are an instance of all the relevant data which exists at the start of the project.
The objectives of data prototyping are to produce:
A set of data cleansing and transformation rules which have been seen to produce data
which is all fit for purpose.
A dataset which is the result of those rules being applied to an instance of the relevant raw
(source) data.
To achieve this, a data architect uses a graphical interface to interactively develop and execute
transformation and cleansing rules using raw data. The resultant data is then evaluated and the rules refined.
Beyond the obvious visual checking of the data on-screen by the data architect, the usual evaluation and
validation approaches are to use Data profiling software[26] and then to insert the resultant data into a test
version of the target application and trial its use.
Prototyping for Human-Computer Interaction
When developing software or digital tools that humans interact with, a prototype is an artifact that is used to
ask and answer a design question. Prototypes provide the means for examining design problems and
evaluating solutions.[27]
HCI practitioners can employ several different types of prototypes:
'Wizard of Oz' prototype: named after the Wizard of Oz (character) in the movie, The
Wizard of Oz (1939 film). This is a prototyping method with which the computer-side of the
interaction is faked by an offsite or hidden human.[28] This prototyping technique is
particularly useful for demonstrating functionality that is difficult or lengthy to engineer, such
as applications like Voice user interface.
role prototype: this prototype may not be engineered or look & feel like a finished product,
but the purpose of this type of prototype is to investigate and evaluation a user need, or what
the prototype could do for the user. They can present features and functionality that the user
might benefit from, to demonstrate what role an artifact like the prototype might fulfill for the
user.[27] A famous example of this kind of prototype would be the block of wood carried by
Jeff Hawkins, when developing the Palm Pilot.
paper prototype: this prototype may use cut paper, cardboard, or other inexpensive
materials to demonstrate an interface. The purpose of this prototype is to test with users,
without having to use a digital tool or develop a program to test functionality.[29] Recently,
paper prototyping has fallen out of favor within certain design circles, particularly because
the low-fidelity nature of this method and the lack of effectiveness when testing with
users.[30]
Scale modeling
In the field of scale modeling (which includes model railroading,
vehicle modeling, airplane modeling, military modeling, etc.), a
prototype is the real-world basis or source for a scale model—such
as the real EMD GP38-2 locomotive—which is the prototype of
Athearn's (among other manufacturers) locomotive model.
Technically, any non-living object can serve as a prototype for a
model, including structures, equipment, and appliances, and so on,
but generally prototypes have come to mean full-size real-world
vehicles including automobiles (the prototype 1957 Chevy has A scale model of an Douglas SB2D
spawned many models), military equipment (such as M4 Shermans, Destroyer in a wind tunnel for testing
a favorite among US Military modelers), railroad equipment, motor
trucks, motorcycles, and space-ships (real-world such as
Apollo/Saturn Vs, or the ISS). As of 2014, basic rapid prototype machines (such as 3D printers) cost about
$2,000, but larger and more precise machines can cost as much as $500,000.[31]
Architecture
In architecture, prototyping refers to either architectural model making (as form of scale modelling) or as
part of aesthetic or material experimentation, such as the Forty Wall House open source material
prototyping centre in Australia.[32][33]
Architects prototype to test ideas structurally, aesthetically and technically. Whether the prototype works or
not is not the primary focus: architectural prototyping is the revelatory process through which the architect
gains insight.[34]
Metrology
In the science and practice of metrology, a prototype is a human-made object that is used as the standard of
measurement of some physical quantity to base all measurement of that physical quantity against.
Sometimes this standard object is called an artifact. In the International System of Units (SI), there remains
no prototype standard since May 20, 2019. Before that date, the last prototype used was the international
prototype of the kilogram, a solid platinum-iridium cylinder kept at the Bureau International des Poids et
Mesures (International Bureau of Weights and Measures) in Sèvres France (a suburb of Paris) that by
definition was the mass of exactly one kilogram. Copies of this prototype are fashioned and issued to many
nations to represent the national standard of the kilogram and are periodically compared to the Paris
prototype. Now the kilogram is redefined in such a way that the Planck constant h is prescribed a value of
exactly 6.626 070 15 × 10−34 joule-second (J⋅s)
Until 1960, the meter was defined by a platinum-iridium prototype bar with two marks on it (that were, by
definition, spaced apart by one meter), the international prototype of the metre, and in 1983 the meter was
redefined to be the distance in free space covered by light in 1/299,792,458 of a second (thus defining the
speed of light to be 299,792,458 meters per second).
Natural sciences
In many sciences, from pathology to taxonomy, prototype refers to a disease, species, etc. which sets a good
example for the whole category. In biology, prototype is the ancestral or primitive form of a species or other
group; an archetype.[35] For example, the Senegal bichir is regarded as the prototypes of its genus,
Polypterus.
See also
3D printing
Clay modeling
Minimum viable product
Rapid prototyping
Test article (disambiguation)
References
1. Blackwell, A. H.; Manar, E., eds. (2015). "Prototype" (http://ic.galegroup.com/ic/scic/Referenc
eDetailsPage/ReferenceDetailsWindow?failOverType=&query=&prodId=SCIC&windowstat
e=normal&contentModules=&display-query=&mode=view&displayGroupName=Reference
&limiter=&currPage=&disableHighlighting=false&displayGroups=&sortBy=&search_within_
results=&p=SCIC&action=e&catId=&activityType=&scanId=&documentId=GALE%7CENKD
ZQ347975681&source=Bookmark&u=dclib_main&jsid=63ed7b4b8ba090d7346c2d926005
d307). UXL Encyclopedia of Science (3rd ed.). Retrieved 13 July 2015.
2. Gero, John S. (1990-12-15). "Design Prototypes: A Knowledge Representation Schema for
Design" (https://www.aaai.org/ojs/index.php/aimagazine/article/view/854). AI Magazine. 11
(4): 26. ISSN 0738-4602 (https://www.worldcat.org/issn/0738-4602).
3. "Prototyping Definition" (https://www.pcmag.com/encyclopedia_term/0,1233,t=prototyping&i
=49886,00.asp). PC Magazine. Retrieved 2012-05-03.
4. Marcelo M. Soares; Francesco Rebelo (15 August 2012). Advances in Usability Evaluation
(https://books.google.com/books?id=Qy_QRu-7dKcC&pg=PA482). CRC Press. p. 482.
ISBN 978-1-4398-7025-9.
5. "prototypical (adjective) definition and synonyms | Macmillan Dictionary" (https://www.macmi
llandictionary.com/dictionary/british/prototypical). www.macmillandictionary.com. Retrieved
2019-12-15.
6. Harper, Douglas. "prototype (n.)" (http://www.etymonline.com/index.php?term=prototype&all
owed_in_frame=0). Online Etymology Dictionary.
7. Lai, Chun Sing; Locatelli, Giorgio (February 2021). "Valuing the option to prototype: A case
study with Generation Integrated Energy Storage" (https://doi.org/10.1016%2Fj.energy.2020.
119290). Energy. 217: 119290. doi:10.1016/j.energy.2020.119290 (https://doi.org/10.1016%
2Fj.energy.2020.119290).
8. "Proof-of-Principle Prototype" (https://3d-printing-expert.com/proof-of-principle-prototype/).
3d-printing-expert.com. Retrieved 2019-12-15.
9. "What Is A Working Prototype" (https://product-design-prototype-experts.com/what-is-a-worki
ng-prototype/). product-design-prototype-experts.com. Retrieved 2019-12-15.
10. "What Is A Visual Prototype" (https://3d-printing-expert.com/what-is-a-visual-prototype/). 3d-
printing-expert.com. Retrieved 2019-12-15.
11. "User Experience Prototype" (https://3d-printing-expert.com/user-experience-prototype/). 3d-
printing-expert.com. Retrieved 2019-12-15.
12. Somiya, Shigeyuki, ed. (2013). Handbook of Advanced Ceramics: Materials, Applications,
Processing, and Properties (https://books.google.com/books?id=tdx7lS4fO9wC&q=form+stu
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g/general-prototypes). ThomasNet. Retrieved 13 July 2015.
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=z-b10XVvN44C). Apress. ISBN 9781430244462.
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ion&i=37675,00.asp). PC Magazine. Retrieved 2012-05-03.
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om/PRODUCTS/applicationsimulationsoftware.htm). www.baseblock.com. Retrieved
2019-12-15.
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uilding-a-learning-organization). Harvard Business Review (July–August 1993). ISSN 0017-
8012 (https://www.worldcat.org/issn/0017-8012). Retrieved 2019-12-15.
23. "5.5 Function Prototypes" (http://h30097.www3.hp.com/docs/base_doc/DOCUMENTATION/
V40F_HTML/AQTLTBTE/DOCU_055.HTM). HP. Retrieved 2012-05-03.
24. Tuch, Alexandre N.; Presslaber, Eva E.; Stöcklin, Markus; Opwis, Klaus; Bargas-Avila,
Javier A. (2012-11-01). "The role of visual complexity and prototypicality regarding first
impression of websites: Working towards understanding aesthetic judgments". International
Journal of Human-Computer Studies. 70 (11): 794–811. doi:10.1016/j.ijhcs.2012.06.003 (htt
ps://doi.org/10.1016%2Fj.ijhcs.2012.06.003). ISSN 1071-5819 (https://www.worldcat.org/iss
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Business Intelligence and Big Data. Lecture Notes in Business Information Processing.
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org/10.1007%2F978-3-319-96655-7_1). ISBN 978-3-319-96655-7.
27. Houde, Stephanie; Hill, Charles (1997-01-01), Helander, Marting G.; Landauer, Thomas K.;
Prabhu, Prasad V. (eds.), "Chapter 16 - What do Prototypes Prototype?" (https://www.scienc
edirect.com/science/article/pii/B9780444818621500820), Handbook of Human-Computer
Interaction (Second Edition), Amsterdam: North-Holland, pp. 367–381, doi:10.1016/b978-
044481862-1.50082-0 (https://doi.org/10.1016%2Fb978-044481862-1.50082-0), ISBN 978-
0-444-81862-1, retrieved 2021-12-13
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rototypes). The Interaction Design Foundation. Retrieved 2021-12-13.
29. Rettig, Marc (April 1994). "Prototyping for tiny fingers" (https://dl.acm.org/doi/10.1145/17527
6.175288). Communications of the ACM. 37 (4): 21–27. doi:10.1145/175276.175288 (https://
doi.org/10.1145%2F175276.175288). ISSN 0001-0782 (https://www.worldcat.org/issn/0001-
0782).
30. Knapp, Jake (2016-01-06). "Paper prototyping is a waste of time" (https://library.gv.com/pape
r-prototyping-is-a-waste-of-time-353076395187). Medium. Retrieved 2021-12-13.
31. "Archived copy" (https://web.archive.org/web/20131028064336/http://www.cfr.org/technology
-and-science/3d-printing-challenges-opportunities-international-relations/p31709). Archived
from the original (http://www.cfr.org/technology-and-science/3d-printing-challenges-opportun
ities-international-relations/p31709) on 2013-10-28. Retrieved 2013-10-30.
32. "Open source architecture lab launched in Tasmania". Architecture News. 2021.
33. Lev, Jiri. "Forty Wall House – 40walls.org" (https://40walls.org/forty-wall-house/). Retrieved
2021-09-29.
34. Burry, Mark (2017). Prototyping for Architects. GB. ISBN 9780500292495.
35. prototype (http://www.collinsdictionary.com/dictionary/english/prototype).
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Retrieved December 07, 2012.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Prototype&oldid=1065660586"
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prototype
Contents
English
Etymology
Pronunciation
Noun
Synonyms
Derived terms
Translations
See also
Verb
Translations
References
Further reading
French
Etymology
Pronunciation
Noun
Derived terms
Further reading
Norwegian Bokmål
Alternative forms
Etymology
Noun
References
Norwegian Nynorsk
Alternative forms
Etymology
Noun
References
English
WOTD – 16 December 2018
Etymology
From French prototype or Late Latin prototypon,[1] from Ancient
Greek πρωτότυπος (prōtótupos, “original; prototype”),[2] from
πρωτο- (prōto-, “prefix meaning ‘first’”) (from πρῶτος (prôtos,
“first; earliest”)) + τῠ́πος (túpos, “blow, pressing; sort, type”)
(from τύπτω (túptō, “to beat, strike”), from Proto-Indo-European
*(s)tewp- (“to push; to stick”)). The word is analysable as proto- + -
type.
Pronunciation
(Received Pronunciation) IPA(key): /ˈpɹəʊtətʌɪp/
(General American) IPA(key): /ˈpɹoʊtəˌtaɪp/, /-ɾə-/
Audio (GA) (file)
Audio (AU) (file)
Hyphenation: pro‧to‧type A prototype (sense 2) of a “lift
system” or flying machine for an
individual person
Noun
prototype (plural prototypes)

1. An original form or object which is a basis for other forms or objects (particularly
manufactured items), or for its generalizations and models. [from late 16th c.]
1658, Thomas Browne, “The Garden of Cyrus. […]. Chapter I.”, in Hydriotaphia, Urne-
buriall, […] Together with The Garden of Cyrus, […], London: […] Hen[ry] Brome […],
OCLC 48702491 (https://worldcat.org/oclc/48702491); reprinted as Hydriotaphia (The English
Replicas), New York, N.Y.: Payson & Clarke Ltd., 1927, OCLC 78413388 (https://worldcat.org/oc
lc/78413388), pages 102–103 (https://archive.org/details/hydriotaphiaurne00browuoft/page/
n131/mode/1up):
And if Jordan were but Jaar Eden, that is, the Riuer of Eden, Geneſar but Ganſar or
the Prince of Gardens; and it could be made out, that the Plain of Jordan were
watered not comparatively, but cauſally, and becauſe it was the Paradiſe of God, as
the Learned Abramas hinteth, he was not far from the Prototype and originall of
Plantations.
1694, [William Sherlock], A Defence of Dr. Sherlock’s Notion of a Trinity in Unity, […],
London: Printed for W. Rogers, […], OCLC 1015514133 (https://worldcat.org/oclc/1015514133),
pages 28–29 (https://books.google.com/books?id=zFw7AQAAMAAJ&pg=PA29):
[T]his Holy Trinity is not Three Divine Attributes, ſuch as Wiſdom, Power, and
Goodneſs; for they are all Three the very ſame with each other, the ſame Wiſdom,
Goodneſs, and Power, and therefore not Three Parts or Attributes of the ſame Deity,
but each is the whole, the Prototype, and its living Image is.
1839 August, “Plautus. [...]”, in Foreign Monthly Review, and Continental Literary
Journal, volume I, number IV, London: D[avid] Nutt, […]; Dulau and Co., […]; Berlin:
Asher; Paris: Gayet and Lebrun, OCLC 47248175 (https://worldcat.org/oclc/47248175), page 417
(https://books.google.com/books?id=gPoEAAAAQAAJ&pg=PA417):
Only one manuscript of Plautus seems to have escaped the general wreck of
ancient literature; and it served as the prototype to all the manuscripts at present
extant.
1872 October 31, “International Metric Commission”, in Nature: A Weekly Illustrated

Journal of Science, volume VI, number 157, London; New York, N.Y.: Macmillan and Co.,
OCLC 64051812 (https://worldcat.org/oclc/64051812), section III (In Reference to the Carrying Out
of the Commission's Decision), article 34, page 544 (https://books.google.com/books?id
=vuRSAAAAcAAJ&pg=PA544), column 2:
The making of the new prototypes of the metre and the kilogramme, the tracing of
the metres, the comparison of the new prototypes with those of the Archives, as
well as the construction of the auxillary apparatus necessary to these operations,
are entrusted to the care of the French section, with the concurrence of the
Permanent Committee, […]
2. An early sample or model built to test a concept or process.
The prototype had loose wires and rough edges, but it worked.
1952 August, “[Washington Reporting] General Dynamics will Build Second Nuclear
Sub”, in The Log, volume 47, number 9, Bristol, Conn.: Miller Freeman Publications,
OCLC 30746570 (https://worldcat.org/oclc/30746570), page 24 (https://books.google.com/books?i
d=AoEPAAAAIAAJ&pg=RA1-PA22), column 3:
General Electric, under contract to the A.E.C., is now building a land-based

prototype of this nuclear-power plant at West Milton, N.Y. A land-based prototype
of the nuclear-power plant for the "Nautilus," developed jointly by the A.E.C.'s
Argonne National Laboratory and the Westinghouse Electric Corporation, is now
being built by Westinghouse, also under contract to the A.E.C.
1961 March, “Talking of trains”, in Trains Illustrated, page 133:
Unfortunately however, what may seem on paper an ideal specification for a

particular type of machine does not always prove to be so in practice and the
German Federal Railway submits prototypes of every new design to long and
exhaustive tests before plans are made to put it into service.
3. (computing) A declaration of a function that specifies the name, return type, and parameters,
but none of the body or actual code.
2005, J. B. Dixit, “Unit-5: Functions and Pointers”, in Sangeeta Dixit, editor,
Fundamentals of Computing, new edition, New Delhi: Laxmi Publications, →ISBN, page
355 (https://books.google.com/books?id=94E3_oJv550C&pg=PA355):
Like any variable in a C program it is necessary to prototype or declare a function

before its use, if it returns a value other than an int. It informs the compiler that the
function would be referenced at a later stage in the program. / For example, / In
program 1, the statement / void display_message(); / is a function prototype or
declaration. Here void specifies that this function does not return any value, and
the empty parenthesis[sic – meaning parentheses] indicate that it takes no parameters
(arguments).
4. (semantics) An instance of a category or a concept that combines its most representative
attributes.
A robin is a prototype of a bird; a penguin is not.
2014, Cecil H. Brown, “A Survey of Category Types in Natural Language”, in S[avas] L.

Tsohatzidis, editor, Meanings and Prototypes: Studies in Linguistic Categorization
(Routledge Library Editions: Linguistics), London; New York, N.Y.: Routledge, →ISBN,
part 1 (On the Content of Prototype Categories: Questions of Word Meaning), page 23 (ht
tps://books.google.com/books?id=k6LIAgAAQBAJ&pg=PA23):
If the robin is the prototype of bird, do particular examples of robin constitute that
prototype for different people? I think not. Rather, prototypes are themselves
categories. Thus, to say that a robin is a prototypic bird is to propose that a class of
similar creatures called robin is a prototype of bird.
2015, [anonymous], “Introduction”, in Words, Affixes, and Clitics as Prototype Categories:

Seminar Paper, [Munich, Bavaria]: GRIN Verlag, →ISBN, page 2 (https://books.google.co
m/books?id=IgaIDQAAQBAJ&pg=PA2):
Although it is common knowledge today that a great many linguistic categories are,
indeed, prototype categories […], a number of linguists still perceive grammatical
categories as being classical in their nature […]. These linguists are reluctant to
accept the idea that prototypicality might be relevant to grammar and that
grammatical categories, like all other categories, can also display prototype
effects.
5. (motor racing) A type of race car, a racing sports car not based on a production car. A 4-
wheeled cockpit-seating car built especially for racing on sports car circuits, that does not
use the silhouette related to a consumer road car.
Synonyms
(basis for other forms or objects): see Thesaurus:exemplar
(motorsport): racing prototype, sports prototype, prototype racecar
Derived terms
(Terms derived from prototype (noun)):
prototypal prototype extension prototypicality
prototype-based prototype theory prototypically
prototype-based prototypic prototyping (noun)
programming prototypical
(motorsport): prototype racing
Translations
original form or object which is a basis for other forms or objects
Arabic: ‫( ﻧﻤﻮذج‬namūḏaj) Interlingua: prototypo

Bulgarian: първообраз (bg) m (pǎrvoobraz), Italian: prototipo (it) m
прототип (bg) m (prototip)
Catalan: prototip (ca) m Japanese: プロトタイプ (ja)
(purototaipu),
Chinese: 原型げんがた
(ja) ( , gengata, げんけい ,
原型雏 genkei)
形樣
Mandarin: (zh) (yuánxíng),
Norwegian:
機
(zh) (chúxíng), (of a machine)
(zh) (yàngjī) Bokmål: prototyp m, prototype m
Nynorsk: prototyp m, prototype m
Czech: prototyp m
Dutch: prototype (nl) n Plautdietsch: Väabilt n
Finnish: prototyyppi (fi) Polish: prototyp (pl) m
French: prototype (fr) m Portuguese: protótipo (pt) m
Galician: prototipo (gl) m Romanian: prototip (ro) n, prim-model n
German: Prototyp (de) m Russian: прототи́ п (ru) m (prototíp),
моде́ль (ru) f (modélʹ), проо́браз (ru) m
Greek: πρωτότυπο (el) n (protótypo)
(proóbraz)
Hebrew: ‫( אב טיפוס‬he) Serbo-Croatian: prȍtotīp (sh) m
Icelandic: frumgerð (is) Spanish: prototipo (es) m
Indonesian: prototipe (id), purwarupa (id) Swedish: prototyp (sv) c
Turkish: prototip (tr), model (tr)
early sample or model built to test a concept or process
Bulgarian: прототип (bg) m (prototip) Polish: prototyp (pl) m

Plautdietsch: Väabilt n
declaration of a function
instance of a category or a concept that combines its most representative attributes
Polish: prototyp (pl) m

motorsports car type
See also
antetype
first article
Verb
prototype (third-person singular simple present prototypes, present participle prototyping, simple past
and past participle prototyped)
1. (transitive) To create a prototype of.
1807 July, Alex. Denmark, T. Bradley and R[obert] Batty, editor, [Mr. Denmark, in Answer
to Mr. Chalmers.] To the Editors of the Medical and Physical Journal., volume XVIII,
number 101, London: Printed for R[ichard] Phillips, by W[illiam] Thorne, […], OCLC
1041709964 (https://worldcat.org/oclc/1041709964), page 66 (https://books.google.com/books?id=
X-waAQAAMAAJ&pg=RA1-PA66):
In short, he has purposely perverted the whole case from beginning to end, and
distorted it in such a manner, as not to be prototyped except by his own mind; […]
1857, Spencer T[imothy] Hall, “The Wye and Dove”, in The Peak and the Plain: Scenes
in Woodland, Field, and Mountain, 2nd edition, London: Houlston and Wright, […], OCLC
559839738 (https://worldcat.org/oclc/559839738), page 345 (https://books.google.com/books?id=
F8GeqGUAngAC&pg=PA345):
[Y]ou may form acquaintance with the Wye before it sees the light, by penetrating
that interesting cavern, Poole's Hole, as I have several times before. It is a
wondrous place, and worthy of a far more dignified name; a sort of crypt in Nature's
vast cathedral; an assemblage of all grotesque, fantastic and beautiful mineral
formations, in a fretted vault not made by man, yet mimicking or prototyping all his
art.
1859, Frederic W[illiam] H[enry] Myers, “Burns Centenary Poems. I.”, in George
Anderson and John Finlay, editors, The Burns Centenary Poems: A Collection of Fifty of
the Best out of Many Hundreds Written on Occasion of the Centenary Celebration, […],
Glasgow: Thomas Murray and Son; Edinburgh: John Menzies; London: Arthur Hall,
Virtue and Co.; Dublin: M'Glashan and Gill, OCLC 315650392 (https://worldcat.org/oclc/31565039
2), stanza V, page 3 (https://books.google.com/books?id=5k0CAAAAQAAJ&pg=PA3):
[W]hatsoe'er the poet sings, / Of prototyped in nature or in man, / Moves deeply,

though it touch not wrath of kings / Or frantic battle-van.
1995, Eugene Fiume, “Mathematical Computation”, in An Introduction to Scientific,

Symbolic, and Graphical Computation, Wellesley, Mass.: A K Peters, →ISBN, section 0.2
(Themes of This Book), page 4 (https://books.google.com/books?id=oYlQAAAAMAAJ&p
g=PA4):
The following themes will arise repeatedly in this book: / • the use of symbolic
computation to prototype the behaviour of models.
2014, James A. Langbridge, “The History of ARM”, in Professional Embedded ARM

Development, Indianapolis, Ind.: Wrox, John Wiley & Sons, →ISBN, part I (ARM Systems
and Development), pages 4–5 (https://books.google.com/books?id=Z29RAgAAQBAJ&p
g=RA1-PA4):
The BBC wanted a computer to go with their television series and started to look
for candidate systems. […] Several companies competed for the contract, and the
Proton project was an ideal candidate. The only problem was the Proton didn't
actually exist. It was only in the design stage; it wasn't prototyped. Acorn had little
time, only 4 days, and spent those 4 days working night and day, prototyping the
design, and getting the Proton ready to show to the BBC. […] The BBC Micro was
born.
Translations
to create a prototype of
References
1. ^ James A. H. Murray [et al.], editors (1884–1928), “Prototype”, in A New English Dictionary
on Historical Principles (Oxford English Dictionary), volume VII (O–P), London: Clarendon
Press, OCLC 15566697 (https://worldcat.org/oclc/15566697), page 1512 (https://archive.org/stream/oe
d07arch#page/1512/mode/1up), column 2.
2. ^ “prototype (https://www.lexico.com/definition/prototype)”, in Lexico, Dictionary.com; Oxford
University Press, 2019–present.
Further reading
prototype on Wikipedia.
French
Etymology
Borrowed from Latin prōtotypus, itself a borrowing from Ancient Greek πρωτότυπος (prōtótupos).
Pronunciation
IPA(key): /pʁɔ.tɔ.tip/
Audio (file)
Noun
prototype m (plural prototypes)
1. prototype
Derived terms
prototypique
Further reading
“prototype (https://www.cnrtl.fr/definition/prototype)” in Trésor de la langue française
informatisé (The Digitized Treasury of the French Language).
Norwegian Bokmål
Alternative forms
prototyp
Etymology
From Ancient Greek πρωτο- (prōto-) + τύπος (túpos)
Noun
prototype m (definite singular prototypen, indefinite plural prototyper, definite plural prototypene)
1. a prototype
References
“prototype” (https://ordbok.uib.no/perl/ordbok.cgi?OPP=prototype&bokmaal=+&ordbok=bok
maal) in The Bokmål Dictionary.
Norwegian Nynorsk
Alternative forms
prototyp
Etymology
From Ancient Greek πρωτο- (prōto-) + τύπος (túpos)
Noun
prototype m (definite singular prototypen, indefinite plural prototypar, definite plural prototypane)
1. a prototype
References
“prototype” (https://ordbok.uib.no/perl/ordbok.cgi?OPP=prototype&nynorsk=+&ordbok=nynor
sk) in The Nynorsk Dictionary.
Retrieved from "https://en.wiktionary.org/w/index.php?title=prototype&oldid=62453428"
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Design and construction of a prototype CMY (Cyan-Magenta-Yellow) genetic circuit

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combination) differences. The evolutionary stability dynamics

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Activity: Visualize and Build a Prototype

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