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The Plant Cell

The Plant Cell

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11/26/2010

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1494 The Plant Cell
LETTER
TO
THE EDITOR
The
Role
of
Auxin in Plant Embryogenesis
In the June 1993 issue of THE PLANTCELL, the research paper by Liu et al.(1993) and the review article by Chasan(1993) addressed the important questionof the role of auxin in plant embryogene-sis. These articles are especiallynoteworthy because hey discuss he liter-ature on both zygotic and somaticembryos, each of which offers distinct ad-vantages for studying the underlyingmechanisms of plant embryogenesis. Theintent of this letter isto place he work ofLiu et al. (1993) in the context of what isalready known about the transport, activ-ity, and biosynthesis of auxin during plantembryogenesis.Polar auxin transport has long beenpostulated to play a central role in plantembryogenesis. Microscale auxin trans-port assays have been used to show thatthe hypocotyls of mature embryos dis-sected from both gymnosperm and an-giosperm seeds exhibit pronouncedpolar auxin transport toward the rootend irrespective of their orientation(Greenwood and Goldsmith, 1970; Fry andWangermann, 1976). This polar transportis completely blocked by 10 VM 2,3,5-triiodobenzoic acid
(TlBA)
(Greenwoodand Goldsmith, lm), which suggests hatthe TIBA-mediated effects on culturedzygotic embryos observed by Liu et al.(1993) are directly attributable to the inhi-bition of polar auxin transport asopposed
to
being a secondary effect unrelated toauxin transport. In addition, Fry andWangermann (1976) were the first to pro-pose that the initiation of polarized auxintransport in the globular embryo mightmediate the morphological polarity ex-pressed in the later stages of plantembryogenesis.The possible role of polar auxin trans-port in somatic embryogenesiswas testedby Schiavone and Cooke (1987), who treat-ed different stages of carrot somaticembryos with TlBA and a different auxintransport inhibitor, N-(l-naphthy1)phtha-lamic acid (NPA). Both auxin transportinhibitors at a concentration of
1
pM
areable to block the ability of somaticembryos to undergo morphogenetic ran-sitions to the subsequent stages: globularembryos undergo persistent spherical ex-pansion, oblong embryos (an intermediatestage n somatic embryogenesis)continueaxis elongation without any cotyledon ni-tiation, and heart embryos developadditional growth axes on their hypocotyls.Further evidence that polar auxin trans-port is regulating somatic embryoformation comes from microsurgical in-vestigations, which have established hatcut pieces of somatic embryos can oftenregenerate shoot structures rom their ap-ical ends and root structures from theirbasal ends (Schiavone and Racusen,1990, 1991). Because the regenera-tion process is sensitive to exogenousauxin as well
as
to transport inhibitors(Schiavone, 1988), it is quite conceivablethat the ability
of
developing somatic em-bryos to maintain structural polarity s
also
dependent on polar auxin transport.
In
contrast to Schiavone and Cooke's(1987) observations that carrot somaticembryos are profoundly affected by po-lar auxin transport inhibitors, Liu et al.(1993) reported hat similarly reated zygot-ic embryos
of
Brassica juncea exhibittypical axis elongation and cotyledon posi-tioning at the embryonic apex, but thecotyledons emerge as a fused collarlikestructure around the apex rather than as
two
discrete lateral structures. Their
in-
terpretation that a reduction n polar auxintransport s responsible or the altered cot-yledon morphology was strengthened bytheir finding that the
pinl-7
mutant ofArabidopsis, which is known to have re-duced auxin transport in its inflorescence(Okada et al., 1991), exhibits he same
col-
larlike cotyledon structure in its embryo.We believe hat the difference n the re-sponses of carrot somatic embryos and
6.
uncea zygotic embryos to auxin polartransport inhibitors suggests a provoca-tive hypothesis about the role
of
auxin
in
embryonic pattern formation.
If
we as-sume that these differences cannot beascribed to species differences or othertrivial factors, then it seems reasonableto hypothesize hat embryonic pattern or-mation is actually regulated by
two
overlapping mechanisms: (1) a positionalmechanism that arises as a maternal ef-fect from the ovular tissue surrounding hezygotic embryo or as a consequence ofthe polarized position of the egg cellandlor early embryo within the embryosac, and (2) an auxin-mediated mecha-nism that is established by the initiationof polar auxin transport in the late globu-lar embryo. Either mechanism is appar-ently sufficient by itself to regulate theinitiation of the embryonic axis and theapical positioning of cotyledon structures,because ransport nhibitos prevent theseprocesses in somatic embryos, whichwould seem to lack an active positionalmechanism, but have no effect on zygot-ic embryos, in which the positionalmechanism appears to persist even
in
em-bryo culture. Finally, the emergence of
two
discrete cotyledons appears to be depen-dent solely on auxin transport becausethe inhibitors block this process in bothsomatic and zygotic embryos. Thishypothesis of overlapping regulatorymechanisms has important experimentalimplications: the positional mechanismcan be manipulated via genetic or physi-ological approaches in zygotic embryosonly, whereas the auxin-mediated mech-anism may be investigated more easily insomatic embryos, which lack the position-al mechanism, than in zygotic embryos.Liu et al. (1993) proposed
two
alterna-tive models of localized auxin synthesisand subsequent transport to explain theorigin of the cotyledons at the end of theglobular stage. Although the available evi-dence on auxin biosynthesis duringsomatic embryogenesis lacks the spatial
 
November 1993 1495
LETTER TO THE EDITOR
resolution needed o discriminate betweenthese two models, it does provide consid-erable insight into how auxin gradientsmay be established in developing em-bryos. Using gas chromatography/massspectrometry, Michalczuk et al. (1992a)demonstrated hat 2P-D-treated embryo-genic callus (whose proembryogenicmasses may be considered the somaticequivalent of preglobular embryos) hasextraordinary levels of endogenous aux-in, i.e., approximately
60
ng/g f. wt. freeand
600
nglg
.
wt.
total
IAA,
which is syn-thesized via a tryptophan-mediatedpathway (Michalczuk et al., 1992b). It isdifficult to imagine how effective auxin gra-dients could be established n preglobularembryos n he presenceof such supraop-timal concentrations.Endogenous
AA
levels in young somat-ic embryos decline precipitously ollowingtransfer to 2,4-D-free medium, with theresult that postglobular embryos containbetween 15 to 30 nglg f. wt. of total
IAA
(Michalczuk
et
al., 1992a), which is lowenough to establish interna1 gradients
of
auxin activity. More important, this
IAA
issynthesized via a different, nontryptophanpathway(Micha1czuketal., 1992b), whichimplies that the induction of this secondpathway is required for the initiation of thepolar auxin transport thought to occur inthe apex of the late globular embryo(Schiavone and Cooke, 1987; Liu et al.,1993). Thus, isolating the genes for thenontryptophan pathway may provide theprobes necessary to discriminate betweenthe two potential sites of auxin biosynthe-sis predicted by the alternative models ofLiu et al. (1993).
Todd J. CookeRichard H. RacusenDepartment
of
BotanyUniversity
of
MarylandCollege Park,
MD
20742
Jerry
D.
CohenHorticultural Crops Quality LaboratoryUSDA Beltsville Agricultura1Research CenterBeltsville, MD
20705
REFERENCESChasan,
R.
(1993). Embryogenesis: New mo-lecular insights. Plant Cell
5,
597-599.
Fry, S.C., and Wangermann,
E.
(1976). Polartransport of auxin through embryos. New
Greenwood, M.S., and Goldsmith, M.H.M.
(1970). Polar transport and accumulation ofindole-3-acetic acid during root regenerationby Pinus lambertiana embryos. Planta 95,Liu,
C.-m.,
Xu,
2.-h., and Chua, N.-H.
(1993).Auxin polar transport is essential for thePhytol.
77,
313-317.297-313.establishment of bilateral symmetry duringearly plant embryogenesis. Plant Cell
5,
621-630.
Michalczuk,
L.,
Cooke, T.J., and Cohen, J.D.
(1992a). Auxin levels at different stages ofcarrot embryogenesis. Phytochemistry
31,
Michalczuk,
L.,
Rlbnicky, D.M., Cooke, T.J.,and Cohen, J.D.
(1992b). Regulation ofindole-3acetic acid biosynthetic pathways ncarrot cell cultures. Plant Physiol. 100,
Okada, K., Ueda,
J.,
Komaki, M.K., Bell, C. J.,and Shimura,
Y.
(1991). Requirement of theauxin polar transport system in early stagesof
Arabidopsis
floral bud formation. Plant Cell
Schiavone, F.M.
(1988). Micromanipulationofsomatic embryos of the domesticated carrotreveals apical control of axis elongationand root regeneration. Development
103,
Schiavone, F.M., and Cooke, T.J.
(1987). Un-usual patterns
of
somatic embryogenesis indomesticated carrot: Developmental effectsof exogenous auxins and auxin transport in-hibitors. Cell. Diff.21,53-62.
Schiavone, F.M., and Racusen, R.H.
(1990).Microsurgery reveals regional capabilities orpattern reestablishment n somatic carrot em-bryos. Dev. Biol.141,211-219.
Schiavone, F.M., and Racusen, R.H.
(1991).Regeneration of the root pole in surgical-ly transected carrot embryos occurs byposition-dependent,proximodistal replace-ment of missing tissues. Development113,1097-1103.1346-1353.
3,
677-684.657-664.1305-1313.

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