FLOWER HEATING     FOLLOWING ANTHESIS              & THE EVOLUTION OF GALL MIDGEPOLLINATION IN SCHISANDRACEAE   AMERICAN J...
Shi Xiao Luo & Dianxiang Zhan     South China Botanical Garden,          Guangzhou, China       Shu-Miaw Chaw      Biodive...
KEY TERMS• Agamospermy -the asexual formation of  embryos and seeds without the occurrence of  fertilization.• Anthesis - ...
•Background Information• Objectives• Materials• Methods• Results• Discussion
BACKGROUND INFORMATION
STAR VINE FAMILY                       ~90 SPECIESSchisandraceae                        Kadsura (22 species) - Endemic to ...
WHY IS THIS PAPER INTERESTING?   • First Data Ever Published On     Thermogenesis & Pollination     In Any Asian Illicium ...
PREVIOUS STUDIES    POLLINATION BIOLOGY       (5 of 90 Species) North American Schisandraceae               &      Asian S...
PREVIOUS EXPERIMENTS SUPPORT:                          FLOWER HEATING:               1.) is a direct energy reward for ect...
NORTH AMERICAN      SPECIES               (New World Illicium & Schisandra)-Pollinated predominantly by flies with beetles...
Illicium floridanum      Jack Scheper - www.Floridata.com
Illicium parviflorum        Photo By:
Schisandra glabra
ASIAN SPECIES                  (Old World Kadsura & Schisandra)-Exclusively Pollinated by pollen-eating, Megomatta gall mi...
Kadsura longipedunculata                           www.plant.ac.cn
Schisandra henryii                     Photo By:
Schisandra henryii   PHOTO BY:
FLOWER HEATING     FOLLOWING ANTHESIS             & THE EVOLUTION OF GALL MIDGEPOLLINATION IN SCHISANDRACEAE
POST ANTHESIS FLOWER HEATINGSchisandraceae        Old World Illicium   EVOLUTION OF GALL MIDGE POLLINATION
OBJECTIVESQUESTIONS AUTHORS ARE TRYING TO ANSWER1.) Given that some Schisandraceae exhibit flower heating, do AsianIlliciu...
MATERIALS
STUDY SPECIES 1: Illicium dunnianumDESCRIPTION: Small shrub (0.5 – 2m high)HABITAT: Riverbanks in wooded ravines (Elevatio...
METHODS
FLORAL DEVELOPMENT, FUNCTION, & TEMPERATURE             Open Pollinated,                         Male Phase,             C...
Plant Reproductive Biology                   -Mating Systems-1.) Randomly selected flowers and marked as controls2.) Flowe...
VISITORS
&
POLLINATORSDiurnal & nocturnal observations made over 150                     hours:              -Recorded Kin...
Molecular Phylogenetics                                      &            Ancestral Trait Reconstruction•    Sequenced com...
RESULTS♦   1st night = Female flowers w/ fully receptive stigmas♦   1st day = interim phase (female to male)♦   2nd night ...
FEMALE PHASE                 (Illicium dunnianum)   Pollination occurred mostly at night   Single midges entered same fl...
MALE PHASE                   (Illicium dunnianum)   When anthers dehisced midges again visited to oviposit   20 eggs wer...
NURSING PHASE                       (Illicium dunnianum)   Onset
of
nursing
phase

chamber
temp.
increased
to
higher
tha...
OBJECTIVES1.   Asian
Illicium
species
do
possess
postanthe:c
flower
hea:ng.2.   The
key
stage
of
thermal
warming
was
well
a...
DISCUSSION‐Flower
hea:ng
evolved
during
the
evolu:on
of
Schisandraceae
and
became
co
opted
in
flowers
pollinated
by
flower
b...
Thank YouFor Your Patience
SOURCES•   http://www.efloras.org/florataxon.aspx?flora_id=610&taxon_id=242326638•
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  • Schisandraceae

    1. 1. FLOWER HEATING FOLLOWING ANTHESIS & THE EVOLUTION OF GALL MIDGEPOLLINATION IN SCHISANDRACEAE AMERICAN JOURNAL OF BOTANY -JULY 2010-
    2. 2. Shi Xiao Luo & Dianxiang Zhan South China Botanical Garden, Guangzhou, China Shu-Miaw Chaw Biodiversity Research Center Taipei, Taiwan Susanne S. Renner Department of Biology, University of Munich Munich, Germany
    3. 3. KEY TERMS• Agamospermy -the asexual formation of embryos and seeds without the occurrence of fertilization.• Anthesis - the period during which a flower is fully open and functional.• Oviposition - to deposit or lay eggs.• Saprophytic - organisms that feed on dead or decaying matter.
    4. 4. •Background Information• Objectives• Materials• Methods• Results• Discussion
    5. 5. BACKGROUND INFORMATION
    6. 6. STAR VINE FAMILY ~90 SPECIESSchisandraceae Kadsura (22 species) - Endemic to Tropical Asia Schisandra (25 species) - Tropical Asia- (Exception - S. glabra)Illicium (42 species) - SE Asia, SE U.S., Mexico, Greater Antilles
    7. 7. WHY IS THIS PAPER INTERESTING? • First Data Ever Published On Thermogenesis & Pollination In Any Asian Illicium Species • TR-71U THERMO RECORDER
    8. 8. PREVIOUS STUDIES POLLINATION BIOLOGY (5 of 90 Species) North American Schisandraceae & Asian Schisandraceae Thien et al., 1983; White and Thien, 1985; Deringer et al.,1999; Liu et al., 2007; Yuan et al., 2007, 2008
    9. 9. PREVIOUS EXPERIMENTS SUPPORT: FLOWER HEATING: 1.) is a direct energy reward for ectothermic pollinators 2.) increases the volatilization of chemicals directed at pollinators3.) helps the plant mimic mammalian feces or carrion attracting saprophytic pollinators 4.) enhances the respiratory release of CO₂, which in combination with other volatile chemicals may stimulate fly oviposition
    10. 10. NORTH AMERICAN SPECIES (New World Illicium & Schisandra)-Pollinated predominantly by flies with beetles as co- pollinators Thien et al., 1983; White and Thien, 1985; Deringer et al.,1999; Liu et al., 2007
    11. 11. Illicium floridanum Jack Scheper - www.Floridata.com
    12. 12. Illicium parviflorum Photo By:
    13. 13. Schisandra glabra
    14. 14. ASIAN SPECIES (Old World Kadsura & Schisandra)-Exclusively Pollinated by pollen-eating, Megomatta gall midges Yuen et al., 2007, 2008
    15. 15. Kadsura longipedunculata www.plant.ac.cn
    16. 16. Schisandra henryii Photo By:
    17. 17. Schisandra henryii PHOTO BY:
    18. 18. FLOWER HEATING FOLLOWING ANTHESIS & THE EVOLUTION OF GALL MIDGEPOLLINATION IN SCHISANDRACEAE
    19. 19. POST ANTHESIS FLOWER HEATINGSchisandraceae Old World Illicium EVOLUTION OF GALL MIDGE POLLINATION
    20. 20. OBJECTIVESQUESTIONS AUTHORS ARE TRYING TO ANSWER1.) Given that some Schisandraceae exhibit flower heating, do AsianIllicium species also possess this trait?2.) Does any flower heating continue after a flower’s sexual functionis over, and if so, what is the adaptive significance of postantheticflower heating?3.) Are Asian Illicium species pollinated by gall midges, or do theyshow “generalized” fly and/or beetle pollination similar to New WorldIllicium?4.) Are flower heating and midge pollination functionally correlated?
    21. 21. MATERIALS
    22. 22. STUDY SPECIES 1: Illicium dunnianumDESCRIPTION: Small shrub (0.5 – 2m high)HABITAT: Riverbanks in wooded ravines (Elevations 300-750m)LOCATION: Shiheqiguan - Nan Kun Shan National Forest Park, ChinaOBSERVED: Mid March to late April – 2008 & 2009# OBSERVED: 53 individuals total: (38 at site A, 8 at site B, 5 at site C)STUDY SPECIES 2: Illicium tsangiiDESCRIPTION: Shrub or small tree (up to 10m)HABITAT: Mixed forests or thickets (Elevations 500-800m)LOCATION: Along road - Nan Kun Shan National Forest Park,ChinaOBSERVED: April to late May – 2008 & 2009# OBSERVED: 56 individuals total
    23. 23. METHODS
    24. 24. FLORAL DEVELOPMENT, FUNCTION, & TEMPERATURE Open Pollinated, Male Phase, Cross Pollinated, Nursing Phase, Bagged Pre-Anthesis Female Phase • Relative position & color of tepals, stamens, and styles. • Presence or absence of secretion. • Timing of; style movements, anther dehiscence, and floral organ wilting/abscission. • Ambient and floral temperatures in open-pollinated flowers recorded for 50 h) using TR-71U thermo recorder (AWESOME!) Readings taken every 5 sec. (male phase, nursing phase, female phase) • One-way ANOVA F-tests and t-tests – using statistical package SPSS • G-test- using Microsoft Excel Poptools 3.0
    25. 25. Plant Reproductive Biology -Mating Systems-1.) Randomly selected flowers and marked as controls2.) Flowers self pollinated from male-phase flower then bagged3.) Flowers bagged to test for agamospermy4.) Flowers cross pollinated then bagged
    26. 26. VISITORS
&
POLLINATORSDiurnal & nocturnal observations made over 150 hours: -Recorded Kinds & Numbers of Visitors -Duration of Visits -Insect Behavior To investigate flowers functional phases • Visitor behavior was monitored inside flowers • Flowers were bagged at end of female phase • Flowers were bagged at end of male phase • Tepal tips were trimmed in male phase flowers • Tepal tips were trimmed at beginning of nursing phase
    27. 27. Molecular Phylogenetics & Ancestral Trait Reconstruction• Sequenced complete internal transcribed spacer (ITS) of ribosomal DNA• Sequenced part of the chloroplast trnL region• Data set comprised 43 species – 2197 aligned nucleotides• Performed Maximum Likelihood analysis under GTR + G model of substitution• Bootstrap support values were estimated w/ 100 replicate heuristic searches
    28. 28. RESULTS♦ 1st night = Female flowers w/ fully receptive stigmas♦ 1st day = interim phase (female to male)♦ 2nd night = flowers entered male phase –last 2-3 nights (anther dehiscence)♦ Day 3-7= Nursing phase begins  Inner tepals close, forming a chamber,  Adaxial filament surfaces and inner tepal bases produce a secretion,  Filaments increased in diameter in control from female to nursing phase  Filaments thickened from female to male and then wilted by nursing.
    29. 29. FEMALE PHASE (Illicium dunnianum) Pollination occurred mostly at night Single midges entered same flower numerous times Only one midge at a time – if occupied other midges would fly away Midge eggs became visible & pollen grains appeared on stigmas after visits Single midges deposited 26 (+/- 3) pollen grains Captured midges carried numerous pollen grains on body New species of Clinodiplosis was likely discovered
    30. 30. MALE PHASE (Illicium dunnianum) When anthers dehisced midges again visited to oviposit 20 eggs were found per flower in the male phase, but flowers with trimmed tepals caused eggs to die The highest number of larvae/ flower were found in the bagged male-phase. Highest number of eggs was found in male phase flowers High temperatures beginning of female phase then dropped to ambient during male Produced little heat during day and throughout male stages
    31. 31. NURSING PHASE (Illicium dunnianum) Onset
of
nursing
phase

chamber
temp.
increased
to
higher
than
in
female
phase
 Midges
did
not
try
to
enter
nursing
phase
flowers Larvae
in
nursing
phase
flowers
were
seen
feeding
on
filament
and
tepal
secre=on.


    32. 32. OBJECTIVES1. Asian
Illicium
species
do
possess
postanthe:c
flower
hea:ng.2. The
key
stage
of
thermal
warming
was
well
aEer
the
sexual
func:on
was
over
in
the
two
 studied
species.

The
only
observable
&
supported
benefit
of
post‐anthe:c
hea:ng
is
 exclusive
to
the
larvae
of
the
pollinator.

The
increase
in
temperature
provides
warm
enough
 temperatures
for
larvae
to
survive
and
the
heat
also
causes
filament
and
tepal
secre:on
 which
the
larvae
feed
on.

The
hea:ng
seems
to
be
a
reward
for
the
pollinator
in
a
plant‐ pollinator
mutualis:c
rela:onship.
3. In
contrast
to
the
New
World
Illicium,
the
two
study
species
Illicium
dunnianum
and
I.
tsangii
 were
pollinated
only
by
Clinodiplosis
gall
midges.
4. Despite
these
findings,
comparison
of
the
ancestral
state
reconstruc:ons
for
two
traits
 showed
that
flower
hea:ng
and
gall
midge
pollina:on
are
not
strictly
correlated.


    33. 33. DISCUSSION‐Flower
hea:ng
evolved
during
the
evolu:on
of
Schisandraceae
and
became
co
opted
in
flowers
pollinated
by
flower
breeding
midges?‐Floral
hea:ng
helps
odor
emission??‐Flower
hea:ng
aids
in
pollen
tube
growth??‐Obligate
outcrossing
may
be
enforced
by
self
incompatability??‐Selec:ve
factors
behind
the
evolu:on
of
flower
hea:ng??‐Are
midges
simply
co‐pollinators??
    34. 34. Thank YouFor Your Patience
    35. 35. SOURCES• http://www.efloras.org/florataxon.aspx?flora_id=610&taxon_id=242326638•

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