BIO 108

2010
Day 7, Lecture 17, Title: Angiosperm life cycle.
Text Readings: Campbell et al. (2008), pgs. 252-253, 600-606, 618-621, 625-634, 801-811.
Topics to cover:
Angiosperm Strategy
Homosporous Life Cycle to Heterosporous Life Cycle
Sporophyte
Flower Structures and Sporangia
Meiosis and Spore Production
Ovule and Female Gametophyte Development
Anther Sac and Male Gametophyte Development
Pollen and Pollination
Double Fertilization makes Zygote and Endosperm
Growth of Endosperm and Sporophytic Embryo
Seeds and Fruits
If Time: Slide Show
Angiosperm Strategy
Gymnosperms and angiosperms are the current living seeded plants... were others in the past
Unlike pines, angiosperms often use animals to help them achieve
pollination and dispersion
so need to interact well with animals... play with their minds?
Why should pre-meds study plants?
Plant's experiments in plant/animal interactions
drugs, poisons, perfumes and foods to attract pollinators, repell herbivores
Therefore plants are a source of drugs: ethnobotany, clinical work, etc...
Use of plants to make other items. Genetically modified crops...
Homosporous Life Cycle to Heterosporous Life Cycle
Review homosporous life cycle; typical of non-seeded plants...
Homosporous, one type of sporangium, makes spore of one size in a sporangium
spore (1N), does mitosis to become gametophyte
gametophyte, via mitosis makes archegonium + antheridium, to make gametes
gametes (1N), egg and sperm, do fertilization to make a zygote
zygote (2N), via mitosis will become a sporophyte
sporophyte (2N), multicellular, in sporangia do meiosis and make spores
contrast to heterosporous gymnosperm and angiosperm (flowering plant) life cycle
Heterosporous, two types of sporangia, makes spores of two distinct sizes
microsporangium, in which meiosis makes microspores
megasporangium, in which meiosis makes megaspores
sporophyte (2N), is the generation that has sporangia and does meiosis

spores (1N), two types, each will do mitosis, and differ in developmental fate
male gametophyte (1N), microgametophyte, does mitosis to make sperm
female gametophyte (1N), megagametophyte, does mitosis to make egg
male and female gametophytes (microgamteophyte and megagametophyte)
in angiosperms the gametophytes are so reduced to so few cells
that can not form complete archegonia and antheridia,
but still make gametes
gametes (1N), egg and sperm meet to do fertilization to make a zygote
zygote (2N), uses mitosis to grow and develop into the sporophyte
embryo (2N), a name for a young sporophyte
Sporophyte
Now will consider how the above fits into flowering plants
Flower Structures and Sporangia
Flower is part of the sporophyte generation
Flower structures
4 whorls of modified leaves: sepals, petals, stamen, carpel
stamen: filament, anthers
carpel (also called pistil): stigma, style, ovary, ovule
complete vs. incomplete flowers: 4 whorls present vs. less than all 4
perfect vs. imperfect flowers: stamen and pistil present vs. at least one lacking
What part of the sporophyte stage of flowering plants does sex?
(If you define "sex" as fertilization then, none...
if you include meiosis as part of sexual life cycle then the sporangia do it...)
What is sexual reproduction?
(fusion of egg and sperm, but also change in ploidy to 1N?)
What do the reproductive structures of animals produce?
(gametes)
Reproductive structures in plants that make the gametes are?
(antheridia and archegonia, or items derived from them, on gametophytes)
Reproductive structures in plants that make spores are? (sporangia on sporophyte)
The text notes the sporophytic reproductive structures, but tends to ignore the
gametophytic reproductive structures... Both are needed for this sexual life cycle.
Meiosis and Spore Production
A sporophyte has sporangia in which it does meiosis to make spores
Sporangia here are: anther sacs, and a region in the ovule
Note megaspore will stay in ovule
Microspore is in anther and may be easily released...
Making spores is a sexual reproductive act.
Note these spores grow into gametophytic plants,
this is reproductive as increasing numbers of individuals
it is NOT asexual reproduction as there is change in ploidy in spore formation
Remember, no gametes yet, so no fertilization, just lots of anticipation...

Ovule and Female Gametophyte Development

(Note: Just as animals have variations on sex, there are variations on how gametophytes develop in
the angiosperms. So what is presented here is the "typical" model.)
The megaspore is held in the ovule, it does mitosis to become a female gametophyte
Again, spore production was a sexual reproductive act
just one that animals do not do...
Consider ovule structures, recall ovule is part of sporophyte and has a sporangium
ovule: funiculus, connects ovule to inner wall of ovary
integuments, wrap around ovule, will later become seed coat
micropyle, opening in integuments, through which pollen tube later enters
nucellus, region of ovule that acts as the megasporangium
Four megaspore nuclei made in one cell, three degrade, one large megaspore left
going for quality here, not quantity, all the cytosol ends up with one nucleus
Development of Female Gametophyte
megaspore does mitosis, mitosis, mitosis = 8 nuclei produced, put in 7 cells
mature female gametophyte has 7 cells: (typically, other versions of this exist...)
antipodals, three cells near to funiculus
synergids, two lateral cells near to micropyle
egg, in between the synergids
central cell, has two nuclei (N+N) called polar nuclei
(recall dikaryotic state of fungi?)
Have we made a gamete? (yes, so this is a sexual organism!)
By mitosis or by meiosis? (mitosis)
notice: protected environment here, haploid so must have viable essential alleles
Note that female gametophyte is held in the ovule, protected by previous generation...
Anther Sac and Male Gametophyte Development
Anther sac is a sporangium, is a part of the sporophyte
makes microspores by meiosis
this is a sexual reproductive act
Microspores do mitosis to become a male gametophyte
Microspore is covered by a cell wall that contains sporopollenin.
Microspore does mitosis, to make two cells
tube cell: will grow into a certain shape...
generative cell: will do mitosis again to generate...
two sperm cells, are actually in the cytosol of the tube cell!
male gametophyte: is also called a pollen grain
this is the organism that will engage in fertilization, as it makes the sperm
Initially stays in the volume of the microspore cell wall, for travelling

Pollen and Pollination

Anther sac ruptures, and releases pollen
pollen grain has waxy covering, including sporopollenin...
Pollen, being just a male, is NOT for dispersal,
Pollen, being a male, has to deliver its sperm to an egg
Pollination is a distinct process, seen in most heterosporous species of plants
Here pollination is movement of pollen from anther sac to stigma
this is not dispersal, have not gotten to a new place where species was not
this is also not fertilization, have not fused gametes yet...
Pollination often involves mind games with animals, such as insects....
nectar or other food, perfumes that act as pheromones,
sepals and petals that look like potential mate...
So pollination is a distinctive process from dispersal and from fertilization...
After pollination
growth of pollen tube down style: some of the fastest known growing cells
chemical signals sent between pollen and sporophytic tissue
gets fed by sporophyte
has to grow to egg in the megagametophyte, in ovule, in ovary...
to deliver sperm to it
Have we made gametes? (yes)
Is this male gametophyte a sexual reproductive organism? (yes)
Double Fertilization makes Zygote and Endosperm
One Egg, Two Sperm.... of such arrangements are afternoon soap operas made...
double fertilization
entry of pollen tube into ovule through the open micropyle
role of synergids to check sperm
sperm put into a synergid cell, into its actual cytosol!
sperm checked to see if right species, and of good "breeding"
If of poor quality, sperm are killed
If of good quality, one sperm (1N) fuses with egg (1N),
other fuses with central cell (N+N)
double fertilization: produces 2N zygote, and 3N endosperm
Growth of Endosperm and Sporophytic Embryo
Being triploid the endosperm grows fast
fills with nutrient rich items in a liquid state to bath growing embryo,
this is the fluid in a coconut...
Endosperm is a dead end, could not do meiosis at all well
So endosperm will be absorbed by the sporpophytic embryo... very altruistic!
Sort of like you absorbing a sibling in the womb to help you grow....

Zygote does mitosis, and becomes a young sporophyte = an embryo, which then grows
endosperm expands first to surround embryo,
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Embryo (2N) forms embryonic structures:

formation of radicle, an embryonic root with a RAM
cotyledon(s), specialized leaves
SAM, will grow new leaves and stems
Note that the young sporophyte is surrounded by tissues of previous sporophyte
this embryo grows in ovule, ovule has integuments as an outer layer
integuments of ovule form seed coat
entire ovule is surrounded by the wall of the ovary for additional protection
Seeds and Fruits
Seed, a mature ovule
Ovule is a part of the sporophytic generation
Ovule is many things at many points in time. It:
is a sporangium, makes a megaspore by meiosis
is site of megagametophyte growth and development,
the sporophyte feeds and protects this female gametophyte
is site of double fertilization, when zygote and endosperm are formed
protects and nurishes the growing embryo
becomes a seed
Seeds can go dormant for thousands of years, a major advantage over spores...
problem of dispersal
homosporous plants disperse spores, these travel on wind well
seed plants have huge seeds, harder to disperse
Fruit
ovary, with ovule(s) in it, matures into a fruit with seed(s) in it
development of ovarian wall becomes the fruit coat
issue of dispersal: various solutions,
disperse by wind, modify fruit coat for dispersion
often involving manipulation of animal behaviors
disperse by animal help
ovarian wall as food, poisons often placed in seeds
or inner ovarian wall forms pit to protect seed
and it passes through the digestive tract of the animal...

If Time: Slide Show

"With these obscene processes and prurient apparitions, the gentle and happy scholar of
flowers has nothing to do."
- John Ruskin, advice to students
not probe too far into stamens and pistils -

Objectives:
Compare and contrast the life cycle of animals with that of flowering plants. Be able to
indicate the ploidy and names of each stage in the life cycles, and to identify the sexual reproductive
structures of flowering plants. Be sure that you can find and understand the roles of each of the
following in the angiosperm life cycle: sporophytes, gametophytes, gametes, spores, sporangia,
zygote, embryo. Be able to contrast homosporous and heterosporous plant life cycles.
Unlike animals where meiosis and fertilization are closely associated in the life cycle, plants
separate these processes of sexual reproduction to different stages of the life cycle. Be sure you
understand how both meiosis and fertilization are sexual and reproductive acts in the plant life cycle.
In flowering plants what structures engage in the meiotic part of sexual reproduction, and what
structures are involved in the fertilization part of sexual reproduction? Describe the process of
double fertilization, including the roles played by the synergid and central cell. Understand the
origin and function of the endosperm in the developing seed.
Be able to describe the parts of a perfectly complete flower and their major functions, versus
that of an imperfectly incomplete flower.
Be able to describe how the angiosperm male gametophyte is formed, what cells it contains
when mature, and the function(s) associated with it in the life of flowering plant species.
Correctly place and identify angiosperm ovule structures and functions it carries out at
various stages of its development from it role as a sporangium to its being a seed.
Compare and contrast the functions of the ovarian wall when it first surrounds immature
ovules, versus later when it surrounds mature seeds, in terms of what the ovarian wall might have to
be doing at each of these different points of the life cycle.
Be able to describe various solutions that plants have evolved to address the problems of
inbreeding, achieving pollination, fertilization, and dispersal of their species.
Figures 30.10, 38.3, 38.5 and 38.7 cover most of the structures and processes described in
this lecture and may be good ones to review.
For review, see self quiz questions #1-3 of chapter 30, and questions #2, 3, and 4 of chapter 38.

Needed overheads and items:

Fig. 30.113e, (Campbell and Reece, 2005) orchid
Fig. 38.03ax1, (Campbell and Reece, 2005) lilly
187_K78_104b_Lilium_floral_bud_ls.jpg
178_K79_104d_Lilium_anthers_xs_late_prophase.jpg
181_K79_104e_Lilium_anthers_xs_metaphase_one.jpg
180_K79_104f_Lilium_anthers_xs_second_meiotic_division.jpg
182_L79_104g_Lilium_anther_xs_tetrads.jpg
183_K79_104h_Lilium_anthers_xs_microspores.jpg
184_K79_104i_Lilium_anthers_xs_pollen.jpg
Fig. 38.6, pollen
185_K81_132_Polygonatum_pollen_tubes_wm.jpg
187_K78_104b_Lilium_floral_bud_ls.jpg
188_K82_104a_Lilium_ovules.jpg
190_K82_104b3_Lilium_megasporocyte.jpg
191_K82_104c2_Lilium_megasporogenesis_anaphase_one.jpg
194_K82_104f3_Lilium_megaspores.jpg
198_K82_104k8_Lilium_mature_megagametophyte.jpg
199_K82_104l_Lilium_ovule_double_fertilization.jpg
157_K60_92a_Capsella_embryo_before_cotyledons.jpg
158_K60_92b_Capsella_embryo_early_cotyledons.jpg
159_K60_92c_Capsella_embryo_bending.jpg
160_K60_92d_Capsella_mature_embryo.jpg
Fig. 38.12, (Campbell and Reece, 2005) development pod
20581_violet_seedpod.jpg
171W3731_witch_hazel-capsules_dehiscence.jpg
Fig. 38.11, dispersion collage
171W3734_peach_drupe_section.jpg
Fig. 38.3, angiosperm gametogenesis

References:
Campbell N.A., J.B.Reece, L.A. Urry, M.L. Cain, S.A. Wasserman, P.V. Minorsky,
R.B.Jackson- 2008-Biology. Eighth edition. Pgs. 252-253, 600-609, 618-621,
625-634, 801-811. Pearson/Benjamin Cummings Press. San Francisco, CA.
Campbell N.A., J.B. Reece- 2005-Biology, 7 th edition. Figures 30.113e, 38.3ax1, 38.12.
Benjamin Cummings Press. San Francisco, CA.

Related issues:
Here are some studies of fertilization as done by flowering plants. One examines the spike in the
cytosolic concentration of Ca +2 in the egg. Others look at the role of synergids in pollen tube
recognition. Others have examined how double fertilization has evolved and why.
Escobar-Restrepo J-M., N. Huck, S. Kessler, V. Gagliardini, J. Gheyselinck, W-C. Yang,
U. Grossniklaus- 2007-The FERONIA receptor-like kinase mediates male-female
interactions during pollen tube reception- Science 317: (#5838, 8/3) 656-660
Faure J-E., C. Dumas- 2001-Fertilization in flowering plants. New approaches for an old
story- Plant Physiology 125: 102-104
Ge L.L., H.Q. Tian, S.D. Russell- 2007-Calcium function and distribution during
fertilization in angiosperms- American Journal of Botany 94: (#6) 1046-1060
Higashiyama T., H. Kuroiwa, S. Kawano, T. Kuroiwa- 2000-Explosive discharge of
pollen tube contents in Torenia fournieri- Plant Physiology 122: 11-13
Kranz E., J. Kumlehn- 1999-Angiosperm fertilization, embryo and endosperm
development- Plant Science 142: 183-197
McCormick S- 2007-Reproductive dialog- Science 317: (#5835, 8/3) 606-607
Double fertilization also occurs in some gymnosperms. Here is an article of one example.
Friedman W.E., J.S. Carmichael- 1996-Double fertilization in gnetales: Implications for
understanding reproductive diversification among seed plants- International
Journal of Plant Science 157: (#6, suppl.) S77-S94
Some angiosperm species have been found to have sex chromosom es but they are not of distinct
sizes. Why that is has been an area of study.
Gerelick R- 2005-Theory for why dioecious plants have equal length sex chromosomesAmerican Journal of Botany- 92: (#6) 979-984
Ming R., J. Wang, P.H. Moore, A.H. Paterson- 2007-Sex chromosomes in flowering
plants- American Journal of Botany 94: (#2) 141-150
In the ovule, megasporogenesis creates a spore, and it grows into a female gametophyte. Studies
of these processes are ongoing.
Eriksen B., M. Fredrikson- 2000-Megagametophyte development in Potentilla nivea
(Rosaceae) form northern Swedish Lapland- American Journal of Botany 87:
(#5) 642-651
Jane W-N- 1997-Ultrastructure of the maturing egg apparatus in Arundo formosana
Hack. (Poaceae)- International Journal of Plant Science 158: (#6) 713-726

How microsporgenesis results in spores and then pollen grains is considered in recent studies.
Also how the pollen tubes grow and how they interact with the sporphytic tissues of the style is of
interest.
Brown R.C., B.E. Lemmon- 2000-The cytoskeleton and polarization during pollen
development in Carex blanda (Cyperaceae)- American Journal of Botany. 87:
(#1) 1-11
Dixit R., J.B. Nasrallah- 2001-Recognizing self in the self-incompatibility response- Plant
Physiology 125: 105-108
Obermeyer G., H. Klaushofer, M. Nagel, M. Hftberger, F.W. Bentrup- 1998-In-vitro
germination and growth of lily pollen tubes is affected by protein phosphatase
inhibitors- Planta 207: (#2) 303-312
Pasonen H-L., P. Pulkkinen, M. Kpyla- 2001-Do pollen donors with fastest-growing
pollen tubes sire the best offspring in an amenophilous tree, Betula pendula
(Betulaceae)?- American Journal of Botany 88: (#5) 854-860
Ressayre A., L. Dreyer, S. Triki-Teurtroy, A. Forchions, S. Nadot- 2005-Post-meiotic
cytokinesis and pollen aperture pattern ontogeny: Comparison of development
in four species differing in aperture pattern- American Journal of Botany 92: (#4)
576-583
Tanaka I., K. Ono, T. Fukuda- 1998-The developmental fate of angiosperm pollen is
associated with a preferential decrease in the level of histone H1 in the vegetative
nucleus- Planta 206: 561-569
Wilhelmi L.K., D. Preuss- 1997-Blazing new trails. Pollen tubes guidance in flowering
plants- Plant Physiol. 113: 307-312
There have been studies of angiosperm sperm cells, and sperm cells in other plant species. There
are reports of plant sperm contributing paternal growth factors that influence pattern formation
during the growth of the embryo.
Bayer M., T. Nawy, C. Giglione, M. Galli, T. Meinnel, W. Lukowitz- 2009-Paternal
control of embryonic patterning in Arabidopsis thaliana- Science 323: (#5930,
3/13) 1485-1488
Mogensen H.L., M.L. Rusche- 2000-Occurrence of plastids in rye (Poaceae) sperm cellsAmerican Journal of Botany 87: (#8) 1189-1192
Southworth D., M. Cresti- 1997-Comparison of flagellated and nonflagellated sperm in
plants- American Journal of Botany 84: (#9) 1301-1311
This study of ovule and pollen growth has truly reached new heights:
Kuang A., M.E. Musgrave, S.W. Matthews, D.B. Cummins, S.C. Tucker- 1995-Pollen
and ovule development in Arabidopsis thaliana under spaceflight conditionsAmerican Journal of Botany 82: (#5) 585-595

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Here is an interesting review about the person who discovered the alternation of generations life
cycle of plants.
Kaplan D.R., T.J. Cooke- 1996-The genius of Wilhelm Hofmeister: The origin of causalanalytical research in plant development- American Journal of Botany 83: (#12)
1647-1660

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