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Lily pads (Nymphaea odorata) exhibit heterophylly where a single plant may have leaves that are submerged, floating, or above (aerial) the surface of the water. Lily pads are placed in a unique situation because each leaf form is exposed to a distinctly different set of mechanical demands. While surface petioles may be loaded in tension under conditions of wind or waves, aerial petioles are loaded in compression because they must support the weight of the lamina. Using standard techniques, we compared the mechanical and morphological properties of both surface and aerial leaf petioles. Structural stiffness (EI) and the second moment of area (I) were higher in aerial petioles, although we detected no differences in other mechanical values (elastic modulus [E], extension ratio, and breaking strength). Morphologically, aerial petioles had a thicker rind, with increased collenchyma tissue and sclereid cell frequency. Aerial petioles also had a larger cross-sectional area and were more elliptical. Thus, subtle changes in the distribution of materials, rather than differences in their makeup, differentiate petiole forms. We suggest that the growth of aerial petioles may be an adaptive response to shading, allowing aerial leaves to rise above a crowded water surface.

The pleiofila phenotype (afaftltl double mutant) of Pisum sativum arises from two single-gene, recessive mutations known to affect the identity of leaf pinnae, afila (af), and acacia (tl). The wild-type leaf consists of proximal leaflets and distal tendrils, whereas the pleiofila leaf consists of branched pinnae terminating in small leaflets. Using morphological measurements, histology, and SEM, we characterized the variation in leaf form along the plant axis, in leaflet anatomy, and in leaf development in embryonic, early postembryonic, and late postembryonic leaves of aftl and wild-type plants. Leaves on aftl plants increase in complexity more rapidly during shoot ontogeny than those on wild-type plants. Leaflets of aftl plants have identical histology to wild-type leaflets although they have smaller and fewer cells. Pinna initiation is acropetal in early postembryonic leaves of aftl plants and in all leaves of wild-type plants, whereas in late postembryonic leaves of aftl plants pinna initiation is bidirectional. Most phenotypic differences between these genotypes can be attributed to differential timing (heterochrony) of major developmental events.

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Vascular plants possess multiple mechanisms for defending themselves against pathogens. One well-characterized defense mechanism is systemic acquired resistance (SAR). In SAR, a plant detects the presence of a pathogen and transmits a signal throughout the plant, inducing changes in the expression of various pathogenesis-related (PR) genes. Once SAR is established, the plant is capable of mounting rapid responses to subsequent pathogen attacks. SAR has been characterized in numerous angiosperm and gymnosperm species; however, despite several pieces of evidence suggesting SAR may also exist in non-vascular plants6–8, its presence in non-vascular plants has not been conclusively demonstrated, in part due to the lack of an appropriate culture system. Here, we describe and use a novel culture system to demonstrate that the moss species Amblystegium serpens does initiate a SAR-like reaction upon inoculation with Pythium irregulare, a common soil-borne oomycete. Infection of A. serpens gametophores by P. irregulare is characterized by localized cytoplasmic shrinkage within 34 h and chlorosis and necrosis within 7 d of inoculation. Within 24 h of a primary inoculation (induction), moss gametophores grown in culture became highly resistant to infection following subsequent inoculation (challenge) by the same pathogen. This increased resistance was a response to the pathogen itself and not to physical wounding. Treatment with β-1,3 glucan, a structural component of oomycete cell walls, was equally effective at triggering SAR. Our results demonstrate, for the first time, that this important defense mechanism exists in a non-vascular plant, and, together with previous studies, suggest that SAR arose prior to the divergence of vascular and non-vascular plants. In addition, this novel moss – pathogen culture system will be valuable for future characterization of the mechanism of SAR in moss, which is necessary for a better understanding of the evolutionary history of SAR in plants.

The leaf blade of Pisum sativum L. possesses proximal leaflets and distal tendrils and is altered by two recessive mutations that affect pinna identity, afila (af) and acacia/tendrilless (tl). Using morphological and histology features we characterized the variation in leaf form and pinna anatomy of the Af and Tl heterozygous genotypes. Our goal was to identify the specific interactions of these genes and the relative influence of each in regulating all structural components of the leaf and pinna phenotypes. The Tl/tl genotypes possess broad tendril - narrow leaflets in the distal region that are histologically more similar to leaflets than to tendrils. The afafTltl leaves have simple tendrils in the distal region and branching complexity that is intermediate between that of aftl and af leaves in the proximal region. Only the most distal tips are slightly expanded. Because the Af and Tl genes interact to affect almost all aspects of leaf and pinna form, they affect development at multiple levels of organization.Key words: acacia, afila, Fabaceae, leaf morphology, leaf anatomy, Pisum sativum.

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