EARLY LEAF ONTOGENY AND SHOOT DEVELOPMENT IN BROWNEA ARIZA (LEGUMINOSAE)

Brownea ariza Benth. (Leguminosae: Caesalpinioideae) shows early shoot tip abortion and subsequent renewal growth from the pseudoterminal bud. This species is unusual in that the entire shoot system is formed before flushing from the bud occurs, shoot tip abortion occurs during flushing, and the aborting portion contains three to six leaves as well as primordial structures varying from hood to peg shape. This study focused on the morphological changes from initiation of scale and foliage leaf primordia in the “resting” renewal bud through bud elongation to flushing and bud abortion. Scanning electron microscopy revealed that embryonic scale leaves are hood-shaped while foliage leaf primordia show early segmentation into leaflets and stipules. No transitional stages were observed. Bud scales and foliage leaves show opposite developmental trends. In bud scales, length at maturity increases from first to last formed, while length decreases in sequentially formed foliage leaves. Early in leaf development the stipules keep pace with the elongation of the rachis. When the bud reaches about one half of its final length the leaf rachis begins to exceed the lengths of its stipules. This young rachis terminates in a distinct mucro that persists until maturity at which time it abscises. Growth patterns indicate that mucro and rachis are a single developmental unit. The early abortion of a shoot tip containing several leaves cannot be easily rationalized. Previous suggestions have involved maintenance of form and ecological adaptation. We add the possibility of elimination of cell progeny encumbered by mutations. From this and other studies of this group, it is clear that at maturity leaves of different species may look alike, e.g., Hymenaea and Colophospermum are bifoliolate; Brownea, Saraca, and others are multifoliolate. However, early stages of leaf ontogeny are quite diverse and may be of systematic value, since these early differences are lost or masked by later development.     

Leaf Development, Metamorphic Heteroblasty and Heterophylly in Berberis s. l. (Berberidaceae)

Shoot development of temperate and tropical members of Berberis s. l. was examined in order to assess: (1) the homology of the spines along the long shoots and the foliage leaves that form on the short shoots; (2) the occurrence of heterophylly and/or heteroblasty in the genus; and (3) the structural correspondence between cataphylls, spines, and foliage leaves. The 1-5-armed spines have been interpreted as modified compound leaves lacking stipules, as a modified lamina (central spine) with stipules (lateral spines), or less often, as transformed branches, or as epidermal outgrowths. On the other hand, the foliage leaves of the short shoots have been interpreted as leaflets of palmately compound leaves. Our results indicate that there are three distinct leaf types per node: (1) Leaves modified in spines spirally arranged in long shoots; (2) foliage, expanded leaves densely arranged in short shoots; and (3) cataphylls protecting axillary buds. The spines are leaf homologs with a clear distinction between the leaf base with stipules, and a laminar portion modified into the 1-5-armed spine; the lateral spines are not stipules as they arise from the marginal meristem of the laminar portion, and not from the leaf base. The foliage leaves also have stipules flanking the leaf base. Both spiny leaves and foliage leaves develop an articulation between the base and the laminar portion. Cataphylls of the short shoots of Berberis s. str. and those of the reproductive short shoots of Mahonia correspond to the entire leaf base, but those of the renewal (vegetative) shoots of Mahonia are spiny and have an odd vestigial pinnately compound lamina. Heterochrony due to ontogenetic truncation caused by the formation of the terminal inflorescence at the apex of the short shoots could be responsible for the lack of petiole/lamina differentiation in the foliage leaves. The spiny long-shoot/foliose short-shoot system of branching in Berberis s. str. appears to be genetically and phylogenetically fixed and not environment-dependent. This represents a clear example of metamorphic heteroblasty sensu Zotz et al. (Botanical Review 77:109–151, 2011) with further occurrence of heterophylly along the short shoots.     

INDETERMINATE GROWTH OF LEAVES IN GUAREA (MELIACEAE): A TWIG ANALOGUE

Leaves of seed plants are generally characterized as organs of determinate growth. In this regard, Guarea and related genera seem unusual in that the pinnately compound leaves of these plants contain a bud at their tip from which new pinnae expand from time to time. Previous studies (based upon superficial examinations of leaf-tip buds) have produced contradictory conclusions regarding how long the leaf apex remains meristematic and produces new pinna primordia. In order to determine whether leaf development in Guarea is truly indeterminate, we microscopically examined leaf-tip buds of G. guidonia and G. glabra. In both species, the leaf apex remains meristematic and continues to produce new pinna primordia as the leaf ages. Unexpanded leaves of G. guidonia contained an average of 23 pinna primordia, while the oldest leaves we examined had initiated an average of 44 total pinnae. In G. glabra, unexpanded leaves contained 8 pinnae, whereas an average of 28 pinnae had been initiated on the oldest leaves. These results indicate that leaf development in Guarea is truly indeterminate. Periodic examination of individual intact leaves indicated that the leaves commonly continue their growth for 2 or more years (observed maximum = 51 months). As new leaflets are initiated at the shoot apex (and subsequently expand in rhythmic flushes), older (basal) leaflets may abscise. In addition, the petiole and rachis of the leaf thicken and become woody as a result of the activity of a vascular cambium. Guarea leaves therefore seem to function as the analogue of a typical twig (stem) in general habit as well as in their indeterminate apical growth and secondary thickening.     

Sorbus cibagouensis sp. nov. (Rosaceae) from Zayü County,southeastern Xizang,China

Sorbus cibagouensis sp. nov. (Rosaceae subfam. Rosaceae), a new taxon from Cibagou National Nature Reserve, Zayü County, southeastern Xizang (Tibet), China, is described and illustrated. It is related to S. monbeigii (Cardot) Balakr., but primarily differs in the number of styles (S. cibagouensis = 5; S. monbeigii = 4) and the shape of stipules and leaves (S. cibagouensis: stipules caducous, small, with entire margin, leaflets in 9–11 pairs; S. monbeigii: stipules persistent, large, serrate, leaflets in 6–8 (–10) pairs).     

Development of leaves and shoot apex protection in Metrodorea and related species (Rutaceae)

Most members of Sapindales are characterized by compound leaves, but several genera also (or only) produce simple or unifoliolate leaves. A few genera may bear stipules or pseudostipules. Little is known about the morphological structure and morphogenesis of these types of leaves in Sapindales, but this information is required for comparative and evolutionary studies. Metrodorea is a Neotropical genus of Rutaceae, comprising species presenting compound and unifoliolate leaves, plus heterophylly, together with an intriguing bud‐protecting structure at the leaf base. The aims of the present study are: (1) to examine leaf morphogenesis in Metrodorea and in closely related species (four Esenbeckia spp., Helietta apiculata and Raulinoa echinata); and (2) to improve our understanding of the morphological evolution of leaves in Metrodorea and Rutaceae. Our data show that the hood‐shaped structure at the base of the leaf in Metrodorea, usually interpreted as a sheath, is, in fact, a pair of united stipules, a synapomorphy of the genus. In the species studied, it is possible to recognize two main types of unifoliolate leaf: early unifoliolate leaves and late unifoliolate leaves. We also found that the number of leaflets in the studied species is dependent on the late or early determination of the leaf primordium, and that loss of leaflets may have been favoured by the restriction of space available for development within the cavity formed by the pair of united stipules. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 267–282.     

STIPULES IN SOME MEMBERS OF THE VITACEAE: RELATING PROCESSES OF DEVELOPMENT TO THE MATURE STRUCTURE

The development of stipules especially their spatial and temporal pattern of initiation in relation to the leaf was investigated in Vitis riparia Michx., cv. Concord, Parthenocissus tricuspidata (Sieb. & Zucc.) Planch., Cissus oblonga (Benth.) Planch., Cissus hypoglauca (F.v.M.) A. Gray, and Cissus rhombifolia Vahl. Early initiation is characterized by the occurrence of a single primordium with a wide insertion on the flank of the shoot apex. Distinguishing between stipule primordia and the leaf primordium is impossible at this early stage. Distinct primordia can only be seen in later stages of development. At maturity, the stipules occupy free lateral positions. Developmental processes such as timing of initiation and zonal growth seem to play an important role in early development. In five of the six taxa examined in this study, the early initiation of stipules, their close association with the leaf and also their faster relative rate of growth during early development appear to give them a characteristic protective function. In contrast, C. rhombifolia stipules are initiated later than the leaf and seem to develop at a slower rate than the leaf proper. Consequently, they never enclose their associated leaf but instead cover the next youngest leaf. Many different criteria are used to distinguish the broad category of stipules, and therefore many interpretations have been made depending on the type of approach that is used. This study attempts to look at stipules in terms of developmental processes and demonstrates a more accommodating leaf/stipule concept which provides a clearer comprehension of the nature of the stipule.     

Reversal of the Direction of Photosynthate Allocation in Urtica dioica L. Plants by Increasing Cytokinin Import into the Shoot

The natural cytokinin import from the root into the shoot of Urtica dioica plants was enhanced by supplying zeatin riboside (ZR) solutions of various concentrations to a portion less than 10 % of the root system after removal of their tips. After 6 h ZR pretreatment of the plants, ¹⁴CO₂ was supplied for 3 h to a mature (source) leaf or to an expanding leaf and the ¹⁴C-distribution in the whole plant was determined after a subsequent dark period of 14 h. ZR substantially increased ¹⁴C fixation by the expanding leaves and also enhanced export of carbon and transport to the shoot apex. The effect of the hormone treatment was, however, more pronounced when the ¹⁴CO₂ was supplied to a mature leaf. In the control plants these leaves exported carbon only to the roots: When the amount of the natural daily ZR input from the roots to the shoot was enhanced by 20%, the bulk of the ¹⁴C exported from a mature leaf moved to the shoot apex and only a minor portion of ¹⁴C was still detected in the root fraction. A several-fold increase of the natural daily ZR input into the shoot resulted in a flow of ¹⁴C only to the growing parts of the shoot. The results suggest control of the sink strength of the shoot apex by ZR in Urtica diocia.     

ABA content in shoots and roots of pea mutants af and tl as related to their growth and morphogenesis

The comparative study of shoot and root growth was carried out, and the level of ABA therein determined in the mutant af and tl and wild-type isogenic lines of pea. The recessive af mutation transformed the leaflets into tendrils, and the tl mutation transformed the tendrils into leaflets. These mutations did not affect the length and number of internodes. In all plants, the level of ABA in the leaves was 3–10 times greater than in the roots, and in the course of vegetative growth it rose in both organs. An increase in the shoot area of tl mutant did not change the dry weight of underground and above-ground parts; therefore, the ratio shoot/root in the mutant was identical to that in the wild-type plants. The maintenance of shoot dry weight in the tl mutant at the level of wild-type plant while its area considerably increased was accounted for by a decrease in the thickness of the leaflet and stipule blades. The level of ABA in the stipules of mutant plants was greater than in the wild-type plants. A decrease in the shoot area in the af mutant brought about a decline in its dry weight; however, the ratio root/shoot was maintained at the wild-type level due to a reduced accumulation of dry weight by the root. The level of ABA in the roots of the af mutant was twice greater than in the leafy forms. ABA was assumed to participate in the control over the root growth exerted by the shoot. The absence of leaflets in the af plants was partially compensated for by expanding stipules. The level of ABA therein was three times higher than in the plants of wild type and comparable with the level in the leaflets of the tl mutant and in the wild-type plants. The role of ABA in the growth and final size of leaf blades is discussed.     

Peltate trichomes in the dormant shoot apex of Metrodorea nigra,a Rutaceae species with rhythmic growth

• In Metrodorea nigra, a Rutaceae species with rhythmic growth, the shoot apex in the dormant stage is enclosed by modified stipules. The young organs are fully covered with peltate secretory trichomes, and these structures remain immersed in a hyaline exudate within a hood-shaped structure. Our study focused on the morpho-functional characterization of the peltate trichomes and cytological events associated with secretion.
• Shoot apices were collected during both dormant and active stages and processed for anatomical, cytochemical and ultrastructural studies.
• Trichomes initiate secretion early on, remain active throughout leaf development, but collapse as the leaves expand; at which time secretory cavities start differentiation in the mesophyll and secretion increases as the leaf reaches full expansion. The subcellular apparatus of the trichome head cells is consistent with hydrophilic and lipophilic secretion. Secretion involves two vesicle types: the smaller vesicles are PATAg-positive (periodic acid/thiocarbohydrazide/silver proteinate) for carbohydrates and the larger ones are PATAg-negative. In the first phase of secretory activity, the vesicles containing polysaccharides discharge their contents through exocytosis with the secretion accumulating beneath the cuticle, which detaches from the cell wall. Later, a massive discharge of lipophilic substances (lipids and terpenes/phenols) results in their accumulation between the wall and cuticle. Release of the secretions occurs throughout the cuticular microchannels.
• Continued protection of the leaves throughout shoot development is ensured by replacement of the collapsed secretory trichomes by oil-secreting cavities. Our findings provide new perspectives for understanding secretion regulation in shoot apices of woody species with rhythmic growth.

     

THE SHOOT APEX OF HYMENAEA COURBARIL (LEGUMINOSAE)

Shoot apices of Hymenaea courbaril are representative of those of many distichous plants, showing sharp intraplastochronic changes as a result of differential growth of the most recently formed primordium and the apical meristem proper. However, an unusual feature of this species is the complex arrangement of stipules. These enclose their own leaves as well as younger structures. Furthermore, the midregions of the stipules fuse ontogenetically at their bases, separating the newly formed leaf from the young shoot. Examination of apices of H. courbaril and H. stilbocarpa with the SEM shows similar morphological features in these two species.     

Die Stipeln derIrvingioideae undRecchioideae und ihre systematische Wertung nebst Bemerkungen zur Holzanatomie und Palynologie

TheSimaroubaceae generally have no true stipules. The stipule-like appendages of some genera proved to be pseudo- or metastipules (Weberling & Leenhouts 1965). There seem to be some exceptions, however: the generaCadellia (incl.Guilfoylia) andRecchia on the one hand, and theIrvingioideae on the other. As these taxa, with exception ofRecchia, have simple leaves, there are no indications that their stipule-like appendages might be pseudo- or metastipules. In regard to their position and ontogeny these appendages behave completely like true stipules. Assuming the view ofForman, one could conceive a morphological line from the long, broadly inserted axillary stipules of mostIrvingioideae to the small scaly triangular stipules ofIxonanthoideae. The similarities between the stipules ofIrvingioideae andErythroxylaceae (already emphasized byHallier and others), become even more evident when their ontogeny is investigated. TheIrvingioideae, therefore, might be regarded as a separate family (perhaps with some relation to theErythroxylaceae,Hallier) or as a subfamily ofIxonanthaceae (Forman).—In addition to data on stipules some results on the palynology and shoot anatomy of the generaCadellia (incl.Guilfoylia) andRecchia are reported. Their relationship with theSimaroubaceae also appears doubtful. If they are to be included, they represent a somewhat isolated group near the base of the family which otherwise has lost its stipules.

Herrn Univ.-Prof. Dr.Walter Leinfellner zum 70. Geburtstag gewidmet.     

Ciclo Di Accrescimento e Differenziazione Delle Gemme in Piante Perenni Nel Territorio Di Bari

Summary

Growth CYCLE AND BUDS DIFFERENTIATION IN PERENNIAL PLANTS GROWING IN Bari AREA. 1) Observations ON Ficus Carica DURING THE YEAR 1946.

The fig tree under observation in Bari started its growth (during the year 1946) in the second week of March and stopped in June second half. The growth period lasted about three months (tab. 1 e 2; diagr. 1).

The most rapid growth occurred during the second week of May (tab. 2; diagr. 1).

From the diagr. 3 it appears that in the fig tree the growth end corresponds at the beginning of the rainless weather; in the meantime the temperature rises gradually (diagr. 4). The atmospheric humidity subsists high also during the summer rainless period (diagr. 5).

Among the four shoots examinated, only one showed a further slow lengthening (13 mm.) from 2nd. August to 6th September. The three others did not grow anymore from June 15 (tab 2; diagr. 1).

The young shoot internodes have different lengths. The basal ones are shorther than the middle ones, which appeared during this speedly growing period (diagr. 2); the last two internodes are very short, so that the nodes are rather contiguous. At each node one leaf unfolds, with two large stipules, which soon fall; on the very last node the stipules do not fall, sheltering the young apical bud, which they completely close in. The corresponding leaflet scarsely grows, falling soon. Neverthless the protection of the two stipules, the first leaflet of the apical bud withers and falls frequently as soon as the bud unfolds at the beginning of spring (fig. 1).

Each leaf axil bears two or three buds (fig. 7). When two buds are present, one is a fruit bud and the second a vegetative one (figg. 11 e 13), when three buds are present, the two lateral ones develop into flowers, while the middle one grows into a vegetative shoot (fig. 12).

The very first sign of the fruit character of a bud is the flattening of the apical bud meristem (fig. 7); successively it deeps into a cuplike body (figg. 8 and 9); on the bottom and along the walls of the cup the female floscules are developing (fig. 10).

The first sign of the vegetative character of a bud is, after the formation of the leaf meristem outline, its evolution with the caratteristically lobed leaves and stipules.

As far as a given height along the shoot (as far as the sixth node in our shoots) the syconia prolong their growth after the shoot lengthening has stopped and ripen completely during August-September (forniti or September-figs). From a certain level ahead, as far as the apical bud (from the seventh to the tenth node in our shoots) the syconia cease their development about contemporarly the vegetative shoot. They are three or four mm. wide and the little flowers are clearly distinguishible inside. So they hang from the nodes as late as next spring time, when they begin to grow again, ripening in June (fioroni or June-figs).

September and June figs then represent a continous acropetal fruit series differentiating at each leaf axil, as soon as the leaflets unfold during the shoot development. We must point out that while the September figs reach the ripe condition through a stopless development in the same year, the June figs stop their growth at a given time, standing a resting period and begin to grow again, ripening the next vegetative season.

As to explain such a behaviour, the presence of an inhibitory stuff is prospected, produced from the more developed figs, which attain a given stadium of development. This stuff would act on those figs, which have not yet reached this particular development stage.

The summer drought checkes the shoot growth and the bud differentiation. The flower buds, which have been too early stopped in their development, are not able to grow again the next season and fall during the apical bud unfolding. Such a condition occurs in the last leaf bud (fig. 4) and in the first leaflet bud inside the vegetative tip (apical bud) (fig. 5). At the beginning of the new vegetation season the shoot awakes with the differentiation of those buds, which did not yet differentiate at the axil of the leaflets inside the apical bud (fig. 6). (See fig. I, where the phenomenon is schematically reproduced).     

Characterization of Hormonal Complex in Pea Phenotypes Differing in Leaf Morphology

Two genotypes of the pea (Pisum sativum L.) with wild-type leaves (variety Orlovchanin, Af/Af genotype) and the afila morphotype (aphyllous variety Nord, af/af genotype) were compared in terms of growth performance and hormonal characteristics of different leaf parts and the whole plant. The replacement of leaflets by tendrils in the afila variety led to a reduction in total dry weight and the area of photosynthesizing surfaces. The loss of leaflets was partly compensated for by rapid expansion of stipules at early stages of plant development and by the hypertrophy of tendrils at later stages. The excessive development of stipules in afila plants was paralleled by the increase in IAA and cytokinin level in their tissues. The hypertrophied development of tendrils and chlorophyll accumulation in tendrils of afila plants was correlated with a high IAA and cytokinin content at a low ABA background level. The elevated content of ABA in tissues of wild-type plants was associated with the preferential development of leaflets and a larger transpiratory surface compared with those in the afila form. It is assumed that this feature ensures the turgescence of wild-type plants. The possible involvement of phytohormones in growth and morphogenesis of pea mutants is discussed.     

Two new endemic species of Ophiorrhiza L. (Rubiaceae: Ophiorrhizeae) from Davao Oriental,Philippines

Two new species of Ophiorrhiza collected from Mt Hamiguitan Range Wildlife Sanctuary in Davao Oriental, Philippines, are herein described and illustrated. Ophiorrhiza erythropilosa is the third Philippine species possessing involucral bracts and is further characterized by its predominantly villose stems that appear red due to strikingly red-violet trichomes, subpersistent bifid stipules, linear involucral bracts without prominent midrib, and 4 mm long urceolate corolla that is villose outside and lightly puberulous inside. Ophiorrhiza hamiguitanensis is characterized by its coriaceous, lanceolate leaves with attenuate apex and base, brochidodromous venation, subpersistent bilobed stipules with slightly recurved acute tips, presence of 1.5–2.5 mm long linear ensiform bracts, heterostylous flowers, clavate calyces and broadly obcordate capsules.     

Shoot apex and spathella: two problematical structures of Podostemaceae–Podostemoideae

The presence of a shoot apex and shoot apical meristem (SAM), said to be absent in subfamily Podostemoideae (Podostemaceae), is confirmed for Marathrum utile and M. foeniculaceum. The vegetative shoot axis is terminated by a small group of meristematic cells which are surrounded by the tissue of the adnate bases of foliage leaves. The slightly bulged tip of the shoot apex is embraced by the youngest leaf, facing the apex with its adaxial side. The study also refers to the spathella, a cup-shaped structure covering obligatorily the young flower bud in Podostemoideae. The occurrence of two separate peaks in the young spathella of M. foeniculaceum supports the view that the spathella is formed by two fused bracts (hypsophylls). The two bracts are perpendicular to the distichous foliage leaves below the spathella. The scaly leaflet on the spathella of A. latifolia apparently does not represent a rudimentary blade of the spathella, but is interpreted as a separate bract. The occasional occurrence of scales below or above the spathella points to a reduction of bracts that were originally present in greater number on the pedicels.     

Stipules inRhodoleia (Hamamelidaceae)

Rhodoleia has long been believed to be the only member of theHamamelidales lacking stipules, and its systematic position has been doubtful. The present investigation shows, in contrast, thatRhodoleia championi Hook. f. produces conspicuous stipules which are, however, restricted to a few leaves of the transition region between bud scales and foliage leaves. In the foliage leaves stipules are apparently reduced, except sometimes in the outermost leaf. The presence of stipules and other correspondences clearly shows thatRhodoleia belongs to theHamamelidaceae.     

A GENETIC SYNDROME AFFECTING LEAF DEVELOPMENT IN PISUM

A recessive foliage mutant of Pisum, designated ‘sinuate leaf’ (sil), was found to have two distinct forms of expression, depending on the background genotype. In an af/af background—wherein leaflets are converted to tendrils—sil/sil plants had adventitious tendrils arising from clefts in the distal portion of the stipule. These adventitious tendrils were morphologically modified, just as were the true tendrils on the same plant, by different allelic combinations at the tl locus. In the standard Af background, sil/sil plants had neither incised stipules nor adventitious tendrils, although they did have undulated and somewhat distorted leaflets and stipules. Because mutant expression was variable in an Af background, classification of segregating populations was uncertain. This uncertainty was removed by taking advantange of pleiotropic effects exerted by sil in the presence of one of the wax mutants, wlo, wb, or wsp. Homozygous wlo plants ordinarily have uniformly waxy stipule surfaces, but when the plants were also homozygous for sil the stipule tips were waxless. Conversely, wb/wb and wsp/wsp plants ordinarily have uniformly waxless stipules, but when wb/wb or wsp/wsp plants were also homozygous recessive for sil the stipule tips were waxy. However, sil had no observable effects of any kind on the stipule tips of plants with stipules reduced in size by the action of st/st. By their individual and combined effects, the foliage mutants used in this study revealed developmental relationships among leaf parts not otherwise evident in non-mutant plants.     

DEVELOPMENT,STRUCTURE AND FUNCTION OF SECRETORY TRICHOMES IN PSYCHOTRIA BACTERIOPHILA (RUBIACEAE)

Mucilage-secreting dendroid trichomes develop from the adaxial epidermis of young stipules surrounding the shoot apex. Each trichome consists of a multicellular stalk from which radiate many branch cells. The trichome has no cuticle and the branch cell walls distally are loose cellulosic frameworks. Dictyosomes produce vesicles whose products are secreted through the plasma-lemma and cell wall. Enlarged portions of the ER are frequently associated with dictyosomes and may be part of the system for synthesis and transport of secretion products. Bacteria, which later occur in leaf nodules, are present in the mucilage surrounding trichomes and young leaves. The latter develop stomata through which the bacteria enter. As stipules and leaves grow out of the apical region, the secretory trichomes degenerate and are replaced by non-secretory ones.     

Weeds of change: Cardamine hirsuta as a new model system for studying dissected leaf development

Cardamine hirsuta, a small crucifer closely related to the model organism Arabidopsis thaliana, offers high genetic tractability and has emerged as a powerful system for studying the genetic basis for diversification of plant form. Contrary to A. thaliana, which has simple leaves, C. hirsuta produces dissected leaves divided into individual units called leaflets. Leaflet formation requires activity of Class I KNOTTED1-like homeodomain (KNOX) proteins, which also promote function of the shoot apical meristem (SAM). In C. hirsuta, KNOX genes are expressed in the leaves whereas in A. thaliana their expression is confined to the SAM, and differences in expression arise through cis-regulatory divergence of KNOX regulation. KNOX activity in C. hirsuta leaves delays the transition from proliferative growth to differentiation thus facilitating the generation of lateral growth axes that give rise to leaflets. These axes reflect the sequential generation of cell division foci across the leaf proximodistal axis in response to auxin activity maxima, which are generated by the PINFORMED1 (PIN1) auxin efflux carriers in a process that resembles organogenesis at the SAM. Delimitation of C. hirsuta leaflets also requires the activity of CUP SHAPED COTYLEDON (CUC) genes, which direct formation of organ boundaries at the SAM. These observations show how species-specific deployment of fundamental shoot development networks may have sculpted simple versus dissected leaf forms. These studies also illustrate how extending developmental genetic studies to morphologically divergent relatives of model organisms can greatly help elucidate the mechanisms underlying the evolution of form.     

A new species of <Emphasis Type="Italic">Chamaecrista</Emphasis> sect. <Emphasis Type="Italic">Chamaecrista</Emphasis> ser. <Emphasis Type="Italic">Flexuosae</Emphasis> (Leguminosae,Caesalpinioideae) from Serra do Cipó, Minas Gerais,Brazil

A new species, Chamaecrista truncata, from southeastern Brazil, is described, illustrated and compared to its putative closest relative, C. parvistipula. The new species belongs to Chamaecrista sect. Chamaecrista ser. Flexuosae which is characterized by asymmetrical leaflets with palmate venation, quadrangular stems and axillary peduncles. Additionally, the venation pattern of the leaflets and the different types of stipules observed within this series are shown.