Review questions for the first hour exam[VERY IMPORTANT]: For every one of the photographs, drawings and diagrams on the lecture web pages, you ought to be able to write what it depicts, label the parts and explain their significance.
Some special embryological words: Learn to talk like a pro.This is a slightly revised version of the vocabulary review page posted on Wednesday, January 25.
Polar bodies: Small cells formed on oocytes by the two meiotic divisions.
Animal pole: The place on an embryo surface where the polar bodies formed.
Vegetal pole: The side of an oocyte or early embryo 180 degrees opposite the animal pole.
Blastomere: Any cell of a cleavage-stage or blastula-stage embryo
Blastula: A stage of development of a sea urchin or an amphibian in which a hollow cavity has formed inside the embryo (with no connection to the outside)
Blastocoel: The hollow (water filled) cavity inside a blastula.
Gastrulation: Active movement of cells from the surface into the interior of a blastula.
"Early gastrula": An embryo in which gastrulation has just begun.
Guess what is meant by "mid-gastrula" and"late-gastrula" stages.
Invagination: folding inward of part of a epithelial sheet of cells
Sea urchin gastrulation occurs by invagination. Human lungs form by invagination of part of the archenteron.
Archenteron: The future digestive tract, formed during gastrulation.
Blastopore: The opening into the archenteron.
In frog and salamander embryos, the blastopore forms below the equator, on the side of the embryo that will become the posterior.
Ingression: Movement of epithelial cells out of an epithelium, and change to becoming mesenchymal. Partly caused by weakening of cell-cell adhesion, and increased adhesion to extracellular fibers on the side of the epithelium toward which the cells move.
For example, the primary and secondary mesenchyme cells of sea urchin embryos are formed by ingression.
In bird, reptile & mammal embryos, gastrulation is by ingression instead of by invagination.
Epithelial fusion Some examples are the fusion of the archenteron with the stomodeum and the fusion of one neural fold with the other neural fold.
Cavitation (Of masses of mesenchymal cells, to form epithelial tubes, or sheets), Reorientation of cells to form an epithelial sheet surrounding a water-filled cavity. The blastocoel is formed by cavitation. Teleost fish form their neural tube by cavitation, but other vertebrates form their neural tubes by folding.
(There is quite a bit of redundancy in the questions below.)
1) Would you expect the first polar body to be haploid, or diploid?
2) What about the ploidy of a second polar body?
3) Suppose that a sperm fused with a second polar body; might it then develop into some part of the person who develops from that oocyte? (explain your reasoning, pro or con.)
4) By what evidence might you be able to detect that part of a person's body developed that way? (hint: You wouldn't actually need to see it happen.)
5) If the spindle of a meiotic division (of an oocyte) got moved away from the animal pole, then the cleavages would divide to oocyte into more equal parts (i.e. either one or both polar bodies would be much bigger, even as big as a quarter or half the oocyte).
What if such abnormal polar bodies developed as parts of the body?
6) Why decide differentiated cell type by such active rearrangements as invagination, ingression and other "morphogenetic movements"?
7) What function is served, do you guess, by keeping extra sets of chromosomes until so late?
8) Is there really a certain "moment of conception" at which time each individual becomes genetically unique?
Incidentally, sea urchin oocytes really do become haploid before fertilization. Also jelly-fish.
9) List 5 important differences between sea urchins versus mammal development.
10) List some important similarities
See the notes from the first lecture (January 11) for the answers to these questions.
11) Fertilization occurs when, relative to meiosis?
[answer: before the second meiotic division is finished]
b) In dog oocytes
c) In Jelly-fish oocytes
d) Name one other phylogenetic group of animals that are like jelly-fish in this respect.
12) What are the three primary germ layers?
hint: E, M, E. 13) What are two organs that develop from each of these, in mammals?
[ectoderm - brain, outer layer of skin
14) Describe at least three active cell rearrangements.
, 15) Sketch cell arrangement in sea urchins at the 4 cell stage.
16) Sketch cell arrangements in a mammal embryo at the 4 cell stage.
17) Do mammal embryos sometimes not have a 4 cell stage? Explain why they might not.
18) Sketch cell arrangements at the blastula and blastocyst stages.
19) Blastocysts are a stage of embryonic development in...
most multicellular animals?
20) Blastulas are a stage of embryonic development in...
most multicellular animals?
21) Morphogenetic cell movements occur during the development of? just sea urchins? etc.
22) Ectoderm, mesoderm and endoderm occur in? just sea urchins? just vertebrates, etc.
23) Hox genes? Are they just in sea urchins, just in vertebrates, or what?
24) Colinearity, of locations of gene transcription versus locations of genes on chromosomes?
25) What is the difference between holoblastic versus meroblastic cleavage?
26) Rapid voltage changes occur in nerve cells, muscle cells and what other kind of cell?
27) If a human oocyte were simultaneously fertilized by two sperm, instead of one, then how many complete sets of chromosomes would it then have?
28) What normally prevents more that one sperm from fertilizing an oocyte?
29) Invent TWO different new methods of contraception based on the normal biological methods that prevent more than one sperm from fertilizing an oocyte.
30) Would it surprise you if hox genes were discovered in Paramecia, or other kinds of one celled animals? Would this change current beliefs in any important way?
31) What effect can mechanical forces sometimes have on undifferentiated stem cells?
Questions marked with a * are not easy, but you will gain a lot from thinking carefully about them. I won't ask many such difficult questions.
Questions marked with ** will not be on the exam.
32) List and briefly describe (which should include drawing a sketch of each) at least four morphogenetic cell movements.
33) Describe and sketch several specific embryological examples of invagination.
The following are nine examples of invagination:
**34) Describe and sketch several specific embryological example of epithelial fusion. Fusion of the archenteron with the stomodeum, in embryos of echinoderms and amphibians. Fusion of the neural folds with each other.
**35) Describe and sketch several specific embryological examples of ingression. In echinoderm embryos, formation of the primary mesenchyme (which secretes the skeleton) and the secondary mesenchyme (which differentiates into muscle cells); formation of the neural crest in vertebrates
**36) Which of these morphogenetic movements consists of epithelial cells becoming converted into mesenchymal cells?
*37) The neural tube of teleosts forms by rearrangement of a solid rod of cells into a hollow epithelial tube. This process is called "cavitation". In what sense is it the reverse of ingression?
38) What is at least one other example in which a solid mass of cells rearranges to form an enclosed cavity?
39) What is a fundamental difference between the blastocyst of mammal embryos and the blastula of echinoderm (sea urchin) embryos?
How similar are they? Make a sketch of each. How could you tell them apart?
40) About what percentage of the cells in a mammal blastocyst will become parts of the body of the animal?
41) What structures are formed by the other 95%of the blastocyst cells?
**42) List at least five examples of embryological changes in shapes or positions of organs in which the driving force was long believed to be growth, in the sense of enlargement and/or mitotic division of cells. Diagrams of six examples are shown in this drawing. Other examples are elongation of capillaries and nerve axons.
*43) Suggest experiments by which you could test whether an embryological process is directly caused by cell enlargement, or is caused by mitosis, or is caused by the exertion of traction by cells.
*44) Sketch from memory the diagram of the branching railway tracks leading to 'brain", "sensory nerves", "skin", "notochord", "dermatome", "myotome", "sclerotome", "kidneys", & "heart".
45) An exam question might consist of this branching pattern, with blanks instead of words, and you would be asked to fill in the blanks.
46) Argue true or false: "The concept of embryonic induction is that the metaphorical railway switches in this diagram are controlled by specific signaling chemicals secreted by nearby cells."
*47) Suggest possible reasons why geometric rearrangements of embryonic cells need to occur in order for the eventual 250 differentiated cell types to be located in the correct anatomical locations, relative to each other. (I am guessing there must be some reason. Why not just form the nervous system by differentiation of cells that were already located in the interior of the blastula, etc.?)
?Perhaps because physical forces are used as signals that contribute to induction of cell differentiation?
?Perhaps migrations of cells cause them to become located next to a sequence of different cells, from which they get a series of different chemical signals, that (for some reason) they need to receive one at a time, in certain sequences?
?Please suggest some other possible reason.
For example, what if the edges of the neural tube failed to fuse with each other?
49) Chemical signals that can stimulate embryonic induction can also cause birth defects: Explain why you could have predicted this.
50) Many scientists now hope to develop methods to re-grow damaged organs, starting with undifferentiated "stem cells". Explain why they will need to use chemicals that can cause embryonic induction.
51) Explain the difference between holoblastic cleavage versus meroblastic cleavage.
52) Compare amounts and distributions of yolk in oocytes of sea urchins, mammals, teleosts, amphibians, and birds.
53) Imagine that you discovered oocytes of some unknown kind of animal, could you predict whether they would undergo holoblastic or meroblastic cleavage, based on size of the oocyte, amount of yolk, and/or un-eveness of yolk distribution? Please explain.
54) Why does mammal gastrulation (i.e. gastrulation of mammal embryos) resemble bird gastrulation much more than mammal gastrulation resembles gastrulation of amphibian embryos?
55) What are the advantages of zebra fish for research? List at least 4.
56) Please rank these advantages in order of importance, and explain your reasons.
57) Are there (Is there?) any advantages that is/are absolute "deal-breakers", in the sense that no animal or plant that lacks that (those) properties could be used as model research organisms?
*58) A certain species of sponge is listed on the internet as being a "model organism", although they do not reproduce in captivity and live only on the Great Barrier Reef, along north-eastern Australia.
*59) Would actual Zebras be as good as Zebra Fish as experimental animals? Why not?
60) Xenopus laevis is now being replaced by Xenopus tropicalis as a "model organism" for the reason that X. laevis turned out to be tetraploid. Figure out why tetrapoidy is such a disadvantage. (Hint: genetic manipulation is more difficult; imagine trying to do gene knockouts, if every gene has 4 copies)
61) Teleost oocytes are spherical until fertilized, but cytoplasm then flows into a hemispherical bump at the animal pole. Suggest what force would be capable of causing this flow? You are welcome to suggest two or more possible forces, or sets of forces.
62) What and where is the "Yolk Syncytial Layer"?
63) What and where is the "Enveloping Layer"
64) What and where are the "Deep Cells"
65) Which of these develop into the body of the fish?
66) Draw cross-sections of teleost embryos at several stages of development?
*67) If you wanted to test whether embryonic induction is a normal part of teleost development, how would you try to do this?
68) How and where do teleost embryos use pinocytosis to change the surface properties of the yolk syncytial layer?
69) By what ingenious method did J.P. Trinkaus greatly increase the visibility of this pinocytosis?
*70) Suppose that a researcher inserted a micropipette through the enveloping layer and sucked out about half of the deep cells (before gastrulation), and then let gastrulation and body formation continue as normally as it could, which of the following might happen?
? A tail-less fish would develop, in which the head and body were normal size, but no posterior organs developed?
? Gastrulation failed to occur, and no parts of the fish developed?
What could each of these results tell us (or hint to us) about normal mechanisms of fish development?
*71) When and where would you expect each of the hox genes to begin being transcribed to make their messenger RNAs? First the most 3-prime hox genes, then the hox gene next to it in the 5-prime direction, then the third one along the 3' - 5' axis?
*72) Design some experiments that would take advantage of one or more of the peculiarities of teleost embryos to help answer some fundamental biological question.
73) Suppose that teleosts have evolved a fundamentally different set of mechanisms to neurulate, gastrulate, or to control cell differentiation (as compared with the mechanisms evolved by mammals) then does that mean that discovery of the human mechanisms will be delayed or even prevented? Argue pro or con.
*74) Deep cells converge toward the body axis equally from right and left, adding themselves to the posterior end of the body (first head, then neck, then,,,, etc and finally tail. What might happen if you physically removed all or most of the deep cells to the left of the body axis, just before they had a chance to add themselves to the body?
And what could each possible result tell you about normal mechanisms of development?
Visualize an experiment in which deep cells from a teleost embryo were inserted into the blastocoel of a gastrulating salamander embryo.
75) Compare this with Spemann and Pröscholdt's discovery of embryonic induction
76) If conjoined twins developed from a teleost embryo, describe and sketch sequential stages by which such twinning could/would occur.
*77) Could conjoined twins develop (in any kind of animal) in such a way that very different combinations of hox genes are transcribed in one twin than the other, at the location where they are merged?
*78) Invent an imaginary sequence of early embryonic events, as different as possible from what actually occurs in teleosts, amphibians, birds or mammals.
*79) What are some testable predictions of the assumption that the reptile&bird pattern of development evolved first in egg-laying species? [I have thought of one! Hint: I got the idea from Ichthyosaurs, which could not crawl up on beaches to lay eggs]
*80) What do primates have in common with rodents that could explain the similarity of implantation of their embryos?
*81) Would it be possible for a species of reptile or bird to have identical twins?
82) Would it be possible for a species of reptile or bird to have conjoined twins?
> 83) If the true answer to one of the preceding questions were proven to be "yes", then what would be the answer to the other question? And explain why?
84) Why can't you observe belly-buttons on birds?
85) If an indentation in the flexible shell of a reptile pushed directly on the anterior end of the primitive streak, then what sort of birth defect might result?
86) Would you expect it to be possible for the same kind of birth defect to occur in a bird?
87) Is it possible for early mammal embryos to implant in the fallopian tubules or other tissues?
88) WHY is it possible for early mammal embryos to implant in the fallopian tubules or other tissues?
89) If a set of genetically identical triplets were born, with two of the three having been inside the same chorion and the third having been surrounded by a completely different chorion, then describe the events that must have happened to produce the bodies of the triplets.
90) If each of a pair of twins were implanted at different parts of a uterus, some distance apart, then what event must have occurred to produce this result?
92) Suppose that a frog was discovered in which part of the body is haploid: What might have gone wrong in meiosis that could have produced this result?
92) Suppose that you had some way to cause development of sperm that have no DNA (or whose DNA has been damaged so badly that it can never replicate or make mRNA).
If you fertilized a fish egg with these sperm, what would be the result?
93) What if, in addition, you prevented the first cleavage division from occurring (in one of your oocytes that were fertilized with inactivated sperm), but then let all further cell divisions occur normally.
94) What are two different ways that you could produce triploid animals?
95) Imagine that an embryo's first polar body was as large enough to become one of the individual embryonic cells, what sort of detectable abnormality could this produce?
*96) Would you tend to guess that conjoined twinning results from
(2) Two (originally separate) embryos adhering together but merging only parts of their bodies.
*97) What kind of evidence would/could help answer this question?
(2) Or would this disprove both possible mechanisms? [Hint: No]
(3) Does the Nobel Prize-winning discovery of Proshöldt and Spemann influence your
conception of what sorts of process might be the cause of conjoined twins?
98) In a few percent of identical twins, one of the twins has its aorta on the right side, and all its abdominal organs (stomach, spleen, intestine, chambers of the heart) are mirror images of the normal geometry of these asymetrical organs. This is called situs inversus viscerum, and can also occur in babies that are not twins. Please invent a possible cause, in general terms, it happens because development of these organs is partly controlled by...?
99) If a pair of twins were conjoined head to back (which doesn't really happen), in what sort of spatial pattern would you expect hox gene expression to occur? Please make a sketch.
100) In sea urchin embryos, the primary mesenchyme cells form which tissue?
101) Primary mesenchyme cells originally develop from cytoplasm located around what location of the oocyte?
102) Please sketch a sea urchin embryo late during the process of gastrulation. On this sketch, label the locations of the blastopore, the blastocoel, the archenteron, the primary mesenchyme, the stomodeum and the animal pole. (Write each of these words, and draw an arrow from the word to the location of each of these structures.)
103) Be able to recognize a sketch of a sea urchin gastrula, label its parts (as in the preceding question), and tell what tissue or organ will develop from each of these parts.
104) Please sketch a frog or salamander embryo early in gastrulation, and label the blastopore, the animal pole and the vegetal pole.
105) Label the parts of the surface of an early amphibian gastrula that will differentiate into nerve cells. (This is not as difficult as it might seem)
106) Sketch an amphibian embryo at the end of gastrulation, and label the locations of the future nerve cells and the blastopore.
107) Sketch an amphibian neurula stage embryo which has previously had some notochord embryo grafted into its blastopore. Label the neural tube, and the second neural tube.
**) Suppose that this same identical effect could be produced by grafting the Hensen's node from a gastrulating bird embryo into the blastocoel of a salamander blastula stage embryo: What result would you predict?
** Suggest possible medical uses for some of those proteins (in the preceding question), in combination with undifferentiated "stem" cells.
108) Please sketch at least four stages of embryonic development of human embryos (Comparable to the mammal embryo sketches on the Jan 11th web page.)
109) List as many differences as you can, between sea urchin development as contrasted with human embryology.
110) Please list as many similarities as you can between sea urchin development and human embryology.
111) Sketch a cross section through the stage of development in which there is the maximum similarity between embryos of all vertebrates (Fish, amphibians, birds, reptiles and humans.
112) Why are human egg cells triploid right after gastrulation?
113) What's the distinction between invagination and ingression? [whether cells remain epithelial or convert to become mesenchymal.
114) In what sense is cavitation the reverse of ingression?
115) Increasing size and density of yolk in oocytes tends to result in gastrulating by ingression rather than invagination. (?) Is that true? What are some examples? Suggest possible reasons.
116) Is there a relation between yolk density and size and holoblastic versus meroblastic cleavage? What is that relation? Can you explain this relation in mechanical terms?
117) Is it accurate to say that (most mammal) embryos cleave like sea urchins but gastrulate like birds? [yes]
If so, what are the reasons for these similarities and differences?
**118) Please sketch a railway branching in the pattern in which vertebrate embryonic tissues subdivide into the three "primary germ layers", and then how each of these subdivides further into brain, etc. etc. notochord etc. etc. lungs. Draw and label as many sub-branches as you can.
*119) What are some kinds of fish that are not teleosts?
120) What are the major advantages of zebra fish for embryological research? (as many advantages as you can remember.)
121) What are the enveloping layer and the yolk syncytial layer? Do they become any particular parts of the fishes bodies? (hint, no; but explain why)
122) What are "deep cells", and what organs of the fish's bodies to deep cells develop into?
123) How is the neurulation of teleost embryos different from the neurulation of other kinds of vertebrates? Would you guess this difference has any relation to the enveloping layer? *HINT: Why might teleosts seldom or never have the birth defect called "spina bifida"?
124) Sketch at least 4 stages of the early embryonic development of frog and salamander embryos.
125) What is a major advantage of frog and salamander embryos for embryological research?
126) What is meant by embryonic induction? What Swedish prize was awarded to Spemann for discovering embryonic induction? Do you think he deserved a prize for tying a human hair around an amphibian egg at the one cell stage?
127) Which UNC professor first proved that embryonic induction occurs in nematode embryos? [Bob Goldstein]
128) (Guess) why many scientists considered that nematodes might probably NOT have any examples of embryonic induction? Hint: It's NOT because they don't have notochords.
129) [Hint] If Driesch had used nematode embryos instead of starfish embryos, why would he have reached different conclusions?
130) What very important fact about normal development was proven by that experiment of tying a hair around the one-cell stage embryo?
131) Why was it important to tie the hair at such an early stage of development, instead of at the blastula stage, for example?
*132) If salami can induce formation of neural tubes, why does that increase the difficulty of finding out what the normal molecular signal is that causes induction in normal embryos (into which nothing has been implanted)?
133) Imagine that extracts of salami could induce differentiation of more bone, wherever you injected the substance, would ten thousand dollars be charged for pieces of salami?
*134) Were those eleven million dollar per researcher "honor"aria paid in return for providing hard work and the deep knowledge needed to find proteins that induce bone formation? Or was it paid to keep the secret how easy it was?
135) Describe the method and result of Wilhelm Roux's famous "hot needle" experiment. What theory was this experiment designed to confirm?
136) In what ways was this experiment and its results a little different from what is usually said or assumed?
*137) What do you think might be the real reasons for the results of this experiment?
**138) Remembering Lewis Wolpert's very popular theory that embryonic regulation is caused by if-not-linear-then-at-least-monotonic "morphogen" chemical gradients, in what sense do the results of Roux's hot needle experiment contradict and maybe even disprove Wolpert's theory? Hint: Would diffusion gradients not be prevented or at least distorted by the hot needle damage to cells? Shouldn't any kind of cell-cell signalling, including induction, have been visibly disturbed by the effects if the hot needles?
*139) Suppose that poking cells with hot needles had stimulated cells to send strong inductive signals and induce nearby undamaged cells to switch from differentiating into skin to differentiate into nerves, instead. After all, if tiny pieces of salami can do it, maybe heat-damaged cells can, also!
Describe what Roux would have observed, if hot needles had released inducing substances. Figure out what his interpretation would probably have been. Would Roux have been the discoverer of embryonic induction, instead of Spemann? Or would such results have been uninterpretable? And would Roux perhaps just not told people about them.
Without being stimulated and guided by some kind of theory, can you either design experiments or interpret their results?
140) If you were going to repeat Roux's hot needle experiment:
i) Why would you NOT use nematodes?
*141) Suppose that, before the discovery of co-linearity, somebody had invented a hypothesis that predicted the existence of hox genes and the geometric correlation between their location on chromosomes and the parts of the body where those genes are selectively transcribed (i.e. if the theory had predicted colinearity), then would scientists have looked for evidence sooner, and would nearly everyone have been convinced that the theory must be the true explanation why colinearity has evolved.
*142) What are possible reasons why some genes change so little in evolution?
143) Describe gastrulation in birds.
144) Describe gastrulation in mammals.
145) In what sense is the first week of human embryology spent building a pre-embryonic structure to house, feed and produce the eventual embryo, where gastrulation, neurulation, somite segmentation etc, can then subsequently occur.
146) Diagram the three different ways that mammals can form identical twins?
*147) Why would you guess that elementary textbooks and Wikipedia assume that identical twins occur by separation of the first two cells?
*148) Invent a fourth possible way that identical twins might have occurred, based on your knowledge of normal early development.
149) Chickens are deliberately "bred" to produce abnormally big yolks, for people to eat. Suggest why a side effect might be an increase in the frequency of identical twinning, including more conjoined twins in those eggs whose yolks are only moderately larger that normal, but separate twins in those with the biggest yolks. (Such phenomena have actually been reported).
150) If it were really true that twins become conjoined because of being pushed together, then could non-identical twins sometimes become conjoined, in addition to identical twins?
*151) Sometimes, one of a pair of identical twins has all his or her heart, stomach and liver arranged and shaped so that they are mirror images of normal hearts, etc. (e.g. The aorta comes out the right side of the heart and the pulmonary artery comes out the left side.) Suggest a hypothesis to explain how and why this happens.
152) What might happen if you pushed two starfish or sea urchin embryos tightly together side by side?
153) What might happen if you killed an embryo of a sea urchin or a frog embryo at the one celled stage, and pushed this dead cell tightly against the side of an undamaged embryo at its one cell stage?
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