More cells, bigger cells or simply reorganization? Alternative mechanisms leading to changed internode architecture under contrasting stress regimes.

Shading and mechanical stress (MS) modulate plant architecture by inducing different developmental pathways. Shading results in increased stem elongation, often reducing whole-plant mechanical stability, while MS inhibits elongation, with a concomitant increase in stability. Here, we examined how these organ-level responses are related to patterns and processes at the cellular level by exposing Impatiens capensis to shading and MS. Shading led to the production of narrower cells along the vertical axis. By contrast, MS led to the production of fewer, smaller and broader cells. These responses to treatments were largely in line with genetic differences found among plants from open and closed canopy sites. Shading- and MS-induced plastic responses in cellular characteristics were negatively correlated: genotypes that were more responsive to shading were less responsive to MS and vice versa. This negative correlation, however, did not scale to mechanical and architectural traits. Our data show how environmental conditions elicit distinctly different associations between characteristics at the cellular level, plant morphology and biomechanics. The evolution of optimal response to different environmental cues may be limited by negative correlations of stress-induced responses at the cellular level.

The point mutation induced by the low-energy N+ ion implantation in impatiens balsamine genome.

To investigate the effect of genome mutations induced by low energy ions implantation in higher plants, genome mutation of Impatiens balsamine mutant induced by low energy N+ ion implantation were analyzed by the RAPD, ISSR and genome sequence. Six out of the 121 ISSR primers and 6 out of the 135 RAPD primers showed that polymorphism ratios between mutants and wild type were 4.96% and 2.89%, respectively. Sequence analysis revealed that base deletions, insertions, and substitutions were observed in the mutant genome comparable to wild type. N+ induced point mutations were mostly base substitution (77.4%), no duplication, long fragments insertions and deletions was found. In all point mutation, adenine (A) was most sensitive to the N+ ion implantation in impatiens. The transition was mainly A --> guanine (G) (15.90%) and thymine (T) --> cytosine (C) (12.55%). Transversion happened in A <--> T (16.74%), which much higher than C <--> G(5.02%), G <--> T(6.69%), A <--> C (7.11%) bases. These findings indicate that low energy ions being a useful mutagen were mostly cause the point mutation in impatiens.

Testing mechanisms and context dependence of costs of plastic shade avoidance responses in Impatiens capensis (Balsaminaceae).

PREMISE OF THE STUDY: Many plant species elongate their shoots in response to neighbor proximity and neighbor height. Although these plastic responses may be beneficial in terms of enhancing light interception, they also may have costs in terms of increased risk of mechanical failure (i.e., lodging or breaking) because of thinner stems. This trade-off between light acquisition and stability may shape the evolution of plastic elongation responses to foliage shade. METHODS: In a field experiment manipulating elongation phenotypes and densities, we tested two hypotheses. We predicted that the risks of mechanical failure depend on plastic elongation and/or on characteristics of the immediate neighborhood, such as density and neighbor height. Further, we predicted that plants that fail mechanically would have reduced fitness. KEY RESULTS: Mechanical failure was earlier and more frequent at high density and showed a complex interaction with neighborhood characteristics such as relative height of the neighbors and the expression of early plasticity. Plants that broke earlier had shorter lifespan and lower reproductive output. CONCLUSIONS: Our results show that depending on the height and density of the group, plastic elongation responses can remain advantageous despite costs of increased risk of mechanical failure of the taller stems, as mechanical failure was not associated with strong costs in terms of reduced lifespan or seed production. The overall benefits of elongation outweigh the costs resulting in selection for elongation at the population level.

Selection on spur shape in Impatiens capensis.

Rapid speciation within some plant families has been attributed to the evolution of floral spurs and to the effect of spur length on plant reproductive success. The flowers of Impatiens capensis (jewelweed) possess a long, curved spur in which nectar is produced and stored. Spur length and curvature varies among plants within one population. Here I document that spur shape is variable in natural populations, variation within plants is less than variation among plants, and spur shape is correlated with components of female and male reproductive success. The apparent natural selection is weakly directional in 1 of 2 years, with greatest seed production and pollen removal occurring in flowers with the greatest spur curvature. Bee pollinator visit length is longest at flowers with highly curved spurs, and they leave less nectar in these spurs than in flowers with straighter spurs. Spur angle evolution may be limited, at least in part, by opposing selection by nectar-robbers who prefer to visit flowers with greater spur curvature. Other factors that might contribute to the maintenance of spur angle variation are temporal variation in the strength of selection and potential genetic correlations of spur shape with other traits under selection.

Partitioning adaptive differentiation across a patchy landscape: shade avoidance traits in impatiens capensis.

Adaptation to different habitat types across a patchy landscape may either arise independently in each patch or occur due to repeated colonization of each patch by the same specialized genotype. We tested whether open- and closed-canopy forms of Impatiens capensis, an herbaceous annual plant of eastern North America, have evolved repeatedly by comparing hierarchical measures of F(ST) estimated from AFLPs to morphological differentiation measured by Q(ST) for five pairs of populations found in open and closed habitats in five New England regions. Morphological differentiation between habitats (Q(HT)) in elongation traits was greater than marker divergence (F(HT)), suggesting adaptive differentiation. Genotypes from open- and closed-canopy habitats differed in shade avoidance traits in several population pairs, whereas patterns of AFLP differentiation suggest this differentiation does not have a single origin. These results suggest that open- and closed-canopy habitats present different selective pressures, but that the outcome of diversifying selection may differ depending on specific closed- and open-canopy habitats and on starting genetic variation. Hierarchical partitioning of F(ST) and Q(ST) makes it possible to distinguish global stabilizing selection on traits across a landscape from diversifying selection between habitat types within regions.

Floral meristem indeterminacy depends on flower position and is facilitated by acarpellate gynoecium development in Impatiens balsamina.

* Floral meristems are generally determinate. Termination of their activity varies with species, occurring after carpel or ovule development, depending on the placentation type. In terminal flowering Impatiens balsamina (cv. Dwarf Bush Flowered) some flowers exhibit meristem indeterminacy; they produce organs from the placenta after ovule development. * Here we provide a detailed description of gynoecium development in this line and explore the basis of the indeterminate nature of some of its floral meristems. * We find that the placenta is sometimes established without complete carpel fusion. Proliferative growth derives from meristematic remnants of the placenta and is more common in the terminal inflorescence. RNA in situ hybridization reveals that IbLFY (Impatiens LFY homologue) is expressed in all meristem states, even in proliferating meristems. Expression of IbAG in axillary flowers is as expected in the meristem, stamens and carpels but absent from the proliferating meristem. * We conclude that I. balsamina has cauline placentation. Incomplete suppression of inflorescence identity in flowers of the terminal inflorescence leads to floral meristem proliferation after ovule development in this species.

Petaloidy and petal identity MADS-box genes in the balsaminoid genera Impatiens and Marcgravia.

Impatiens and Marcgravia have striking morphological innovations associated with the flowers. One of the sepals in Impatiens is spurred and petaloid, while in Marcgravia the petals are fused into a cap and nectary cups are associated with the inflorescence. Balsaminaceae (Impatiens) and Marcgraviaceae have surprisingly been shown to be closely related, since both belong to the balsaminoid clade of Ericales (basal asterids). However, several thorough morphological studies thus far have not revealed shared derived characters (synapomorphies) that support a close relationship between these families. In the balsaminoid clade, transitions from entirely green flowers to flowers with heterotopic petaloid organs can be observed. The primary role of class B genes in core eudicots is to specify the identity of petal and stamen floral organs. E-class genes, of which SEP3 is a representative, have been identified as redundant mediators that confer transcriptional activation potential on protein complexes that specify organ identity. Given the conserved function of organ-identity MADS-box genes in model plants, but the rapid molecular evolution in angiosperms, it remains controversial whether these genes have been involved in shaping floral diversity. We have identified a SEP3-like gene and a total of five class B genes from Impatiens hawkeri and Marcgravia umbellata and report their quantitative expression in the floral organs. In Impatiens, two AP3/DEF-like genes were identified with strongly divergent C-terminal domains, one truncated and one unusually long. Both genes show a gradual decrease in expression towards the outer perianth organs, but no GLO-like gene expression is observed in the petaloid sepal. Remarkably, SEP3-like gene expression in the Impatiens perianth is absent from the green sepals but present in the petaloid sepal and in the petals. Dimeric protein interactions of the cloned Impatiens genes were studied in yeast and by using gel retardation. In Marcgravia, strong overlapping class B gene expression is limited to the stamens, but a SEP3-like gene is strongly expressed in the Marcgravia nectary, indicating that both Impatiens and Marcgravia show heterotopic expression of a SEP3-like gene. We discuss several candidate mechanisms for heterotopic petaloidy involving modified gene expression and protein interaction of SEP3-like and class B genes.

Herbivory and competition interact to affect reproductive traits and mating system expression in Impatiens capensis.

As a step toward understanding how community context shapes mating system evolution, we investigated the combined role of two plant antagonisms, vegetative herbivory and intraspecific competition, for reproduction and mating system expression (relative production of selfing, cleistogamous and facultatively outcrossing, chasmogamous flowers and fruits) of Impatiens capensis. In a survey of I. capensis populations, we found that vegetative herbivory and intraspecific competition were positively correlated. In a greenhouse experiment where leaf damage and plant density were manipulated, multispecies interactions had dramatic effects on reproductive and mating system traits. Despite having additive effects on growth, herbivory and competition had nonadditive effects for mating system expression, chasmogamous fruit production, flower number and size, and cleistogamous flower production. Our results demonstrate that competitive interactions influence the effect of herbivory (and vice versa) on fitness components and mating system, and thus antagonisms may have unforeseen consequences for mating system evolution, population genetic diversity, and persistence.

LEAFY, TERMINAL FLOWER1 and AGAMOUS are functionally conserved but do not regulate terminal flowering and floral determinacy in Impatiens balsamina.

In Impatiens balsamina a lack of commitment of the meristem during floral development leads to the continuous requirement for a leaf-derived floral signal. In the absence of this signal the meristem reverts to leaf production. Current models for Arabidopsis state that LEAFY (LFY) is central to the integration of floral signals and regulates flowering partly via interactions with TERMINAL FLOWER1 (TFL1) and AGAMOUS (AG). Here we describe Impatiens homologues of LFY, TFL1 and AG (IbLFY, IbTFL1 and IbAG) that are highly conserved at a sequence level and demonstrate homologous functions when expressed ectopically in transgenic Arabidopsis. We relate the expression patterns of IbTFL1 and IbAG to the control of terminal flowering and floral determinacy in Impatiens. IbTFL1 is involved in controlling the phase of the axillary meristems and is expressed in axillary shoots and axillary meristems which produce inflorescences, but not in axillary flowers. It is not involved in maintaining the terminal meristem in either an inflorescence or indeterminate state. Terminal flowering in Impatiens appears therefore to be controlled by a pathway that uses a different integration system than that regulating the development of axillary flowers and branches. The pattern of ovule production in Impatiens requires the meristem to be maintained after the production of carpels. Consistent with this morphological feature IbAG appears to specify stamen and carpel identity, but is not sufficient to specify meristem determinacy in Impatiens.