Re-interpretation of Wexfordia hookense from the Upper Devonian of Ireland as an arborescent lycophyte

New collections of pyritized axes of the lycophyte Wexfordia hookense have been made from the Upper Devonian (uppermost Famennian) type locality at Sandeel Bay, County Wexford, in south-eastern Ireland. The specimens reveal additional histological features that permit reinterpretation of the morphology of this taxon and reevaluation of its taxonomic affinities. Wexfordia is shown to possess both secondary xylem, with narrow, uni- to biseriate rays, and periderm. The range of variation in relative amounts of primary and secondary xylem can be correlated with position in the mature plant. This evidence indicates that Wexfordia was a small tree rather than an herbaceous form. Fine structure of tracheids and additional anatomical features strongly support affinities with Carboniferous arborescent Isoetales, rather than Devonian Protolepidodendrales, and further support the hypothesis that radiation in this lineage was well underway prior to the Carboniferous.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 275–287.     

Belowground rhizomes and roots in waterlogged paleosols: Examples from the Middle Jurassic of Beijing,China

Plant rhizomes and roots occur in terrestrial ecosystems since at least the Devonian, but the documentation of belowground plant tissues is sparse in the fossil record. In this study, fossils representing belowground rhizomes and roots are described from the top of the Upper Yaopo Formation (Middle Jurassic), at the Yuejiapo section, Mentougou District, Beijing, China. Morphological studies of the plant fossils, together with lithofacies analyses, provide new information on plant–soil interactions during the Jurassic period. Three types of rooting systems are recognized from two fossiliferous beds. The Bed-1 Flora is interpreted as representing a Cladophlebis-dominated community, where abundant foliage remains mainly of Cladophlebis cf. scariosa and Cladophlebis delicatula are associated with Type-A rooting system. The Bed-2 Flora includes Type-B and Type-C rooting systems, although the floristic composition is unknown due to the absence of identifiable foliage remains. The Type-A and Type-B rooting systems consist of abundant in situ vertical rhizomes, fine shoot-borne roots and lateral roots, and are consistent with those of some extant ferns. The Type-C rooting system shows a thick central taproot and at least three orders of lateral roots, an architecture typical of various gymnosperms. The in situ rooting systems, as well as sedimentary evidence, contribute to the recognition of stacked, reworked Entisols in a dynamic waterlogged environment.     

Microconchid encrusters colonizing land plants: the earliest North American record from the Early Devonian of Wyoming,USA

Caruso, JA. & Tomescu, AM.F. 2012: Microconchid encrusters colonizing land plants: the earliest North American record from the Early Devonian of Wyoming, USA. Lethaia, Vol. 45, pp. 490–494. Plant fossils in the Early Devonian Beartooth Butte Formation (Wyoming, USA) are colonized by microconchid encrusters which are found on several plant taxa, at two fossil localities in the formation, and whose tube coil diameters range from 230 to 1170 μm. Colonization is densest on broad Drepanophycus devonicus stems where microconchid individuals encompassing broad size ranges co‐occur in close vicinity. This indicates exposure to microconchid colonization and, therefore, submergence of the plant material for relatively extended periods of time prior to burial. For in situ preserved Drepanophycus, this suggests that the plants grew partially submerged and their submerged parts were colonized by microconchids while still alive. In turn, this indicates that by the Early Devonian microconchids were colonizing freshwater environments. The Beartooth Butte Formation provides the first record of plant colonization by microconchids in North America and, along with only one other Early Devonian record from Germany, the oldest evidence for microconchids colonizing plant substrates. □Devonian, encrusters, microconchid, vascular plants, Wyoming.     

Investigating Devonian trees as geo‐engineers of past climates: linking palaeosols to palaeobotany and experimental geobiology

We present the rationale for a cross‐disciplinary investigation addressing the ‘Devonian plant hypothesis’ which proposes that the evolutionary appearance of trees with deep, complex rooting systems represents one of the major biotic feedbacks on geochemical carbon cycling during the Phanerozoic. According to this hypothesis, trees have dramatically enhanced mineral weathering driving an increased flux of Ca²⁺ to the oceans and, ultimately, a 90% decline in atmospheric CO₂ levels through the Palaeozoic. Furthermore, experimental studies indicate a key role for arbuscular mycorrhizal fungi in soil–plant processes and especially in unlocking the limiting nutrient phosphorus in soil via Ca‐phosphate dissolution mineral weathering. This suggests co‐evolution of roots and symbiotic fungi since the Early Devonian could well have triggered positive feedbacks on weathering rates whereby root–fungal P release supports higher biomass forested ecosystems. Long‐standing areas of uncertainty in this paradigm include the following: (1) limited fossil record documenting the origin and timeline of the evolution of tree‐sized plants through the Devonian; and (2) the effects of the evolutionary advance of trees and their in situ rooting structures on palaeosol geochemistry. We are addressing these issues by integrating palaeobotanical studies with geochemical and mineralogical analyses of palaeosol sequences at selected sites across eastern North America with a particular focus on drill cores from Middle Devonian forests in Greene County, New York State.     

The herbaceous lycophyte Haskinsia Grierson and Banks from the Devonian of western Venezuela, with observations on leaf morphology and fertile specimens

The Devonian herbaceous lycophyte genus Haskinsia Grierson and Banks is characterized by narrow dichotomizing stems and helically arranged petiolate leaves with a broad, approximately deltoid, lamina. New specimens are reported from four horizons in the Lower Member of the Campo Chico Formation, Sierra de Perijá, western Venezuela. H. sagittaka Edwards and Benedetto has sagittate leaves, and fertile specimens are reported for the first time; the ellipsoidal/obovoid adaxially-flattened sporangium is attached to unmodified leaves towards the top of the petiole in the adaxial position. H. sp. has sagittate leaves which are larger than any yet attributed to the genus but is represented by only three specimens. H. hastata sp. nov. has hastate leaves; the sporangia are more globose but positioned similarly to those of H. sagittata. The occurrence of the sporangia on unmodified microphylls indicates that Haskinsia is a member of Protolepidodendrales. The status of Haskinsiaceae is discussed. The Venezuelan specimens are compared with other records of Haskimi and Haskinsia -like plants from North America, Kazakhstan, North China, North Africa and Antarctica.     

Effects of MAPKK inhibitor PD98059 on the gravitropism of primary roots of maize

Gravity plays a fundamental role in plant growth and development, yet the molecular details of gravitropism is not fully understood. Here, we report the effects of PD98059, a specific inhibitor of mitogen-activated protein (MAP) kinase kinase, on the gravitropism of primary roots of maize. Unilateral application of PD98059 to horizontal roots led to different gravitropic growth. Placing PD98059-containing agar on the upper side of the root tips accelerated gravitropic curvature, whereas placing the agar on the lower side inhibited gravitropic curvature. However, no effect was detected when asymmetric application of PD98059 to vertical roots. Global application of maize primary root with PD98059 suppressed root gravitropism. Furthermore, the effects of H₂O₂ on horizontal root gravitropism and vertical root bending were compromised by pretreatment with PD98059. These results suggest an involvement of MAP kinase pathway(s) in gravitropism of maize roots.     

Herbaceous eudicot Fairlingtonia from the Lower Cretaceous of Jiuquan Basin,Northwest China and its radiation in Laurasia

Eudicots exhibit diverse life forms and occupy a wide variety of habitats in the modern terrestrial ecosystems, and the diversification began during the Early Cretaceous; however, few Early Cretaceous fossils are preserved as multiorgan whole plants that can provide sufficient morphological characters for detailed phylogenetic assessment. Here, Fairlingtonia microgyna sp. nov. is reported from the upper Lower Cretaceous of Zhonggou Formation, Hanxia Section, Yumen City, western Gansu Province, Northwest China. The specimen is exceptionally preserved as multiorgan whole plant fossil with fibrous adventitious roots, simple and deeply dissected leaves, solitary and dehiscent capsular fruits attached to the creeping stems. As such, it was interpreted as a herbaceous eudicot. Phylogenetic analyses support a placement within the Papaveraceae, most likely in Papaveroideae, but there are obvious differences in morphological characteristics, which cannot confirm the systematic position within the Papaveraceae. Fossil records of Fairlingtonia from contemporaneous deposits (late Aptian to early Albian) in Northwest China and eastern North America provide direct evidence of the geographical radiation of Fairlingtonia on Laurasia. And the morphological characters of F. microgyna, including creeping leafy branches, fibrous adventitious roots, small and deeply dissected leaves as well as small capsular fruits with tiny seeds probably indicate that it was a colonizer of lake-shore environments under wet and bright conditions and possessed fast-growing and rapid propagation habitats, which allowed it to expand its geographic range with both sexual and asexual reproduction.     

Tree fern growth strategy in the Late Devonian cladoxylopsid species Pietzschia levis from the study of its stem and root system

Portions of stems from five new anatomically preserved specimens of Pietzschia levis from a new Late Devonian plant locality of eastern Tafilalt, Anti-Atlas (Morocco), were analyzed to complete the preliminary reconstruction previously done with a single specimen. The basal part of the longest new specimen consists of an obconical portion of stem surrounded by a thick mantle of adventitious roots. Roots are connected to the peripheral strands of primary xylem specific to the stele of Pietzschia stems. Roots grow outwardly; they cross the cortex and the broad central pith at a steep angle and emerge from the stem lower down. The number of roots produced at one level increases conspicuously from the base towards the distal end of the obconical portion of stem. By contrast, cross-sectional dimensions of roots at their origin level decrease distally. Individual roots increase in diameter, and their stele gets more lobed as they grow through stem tissues. The large number of roots at the specimen base and their wider dimensions at this level contribute to the conspicuous enlargement of the stem base. Patterns assessed from the reconstruction of the Pietzschia levis root system may be close to those of the older cladoxylopsids Pseudosporochnales comprising an upright trunk. Growth strategies in the small-statured species P. levis and in younger arborescent ferns of the Psaronius type are compared. They differ mainly in the relative lengths of epidogenetic vs. apoxogenetic growth phases of the stem.     

Root architecture,early development and basal sweep in containerized and bare-rooted Douglas fir (Pseudotsuga menziesii)

Seedlings grown in containers often have deformed, spiralling, kinking root systems (reported especially in pine growing in colder climates like Fenno-Scandinavia). The current study examines the effect of containers on the root systems of Douglas fir planted in Ireland. A sample of sixteen 10-year-old Douglas fir trees, planted as either bare-rooted transplants (2/1) or containerized seedlings (Paperpot 610 (2/0)) on an acid brown earth site in Ireland, were excavated for root achitectural studies. In addition to thorough above-ground measurements, an assessment of basal sweep was also carried out. Root systems were systematically described and horizontal angles, cross-sectional areas (CSA), and maximum depths of all roots>5 mm in diameter were recorded. Various biomass ratios and estimates for dominant roots were also calculated. Symmetry of the root systems, max. and min. numbers, and CSA of roots for sectors of various sizes were compared for the two plant types. The initially (when planted) bigger bare-rooted transplants were still larger than the container-planted trees after ten years. Of the total above-ground fresh weight, the container stock had allocated more to the crown and less to the stem compared to bare-roots. The difference in root weights between stock types was small. Basal sweep occurred on average in 50% of the bare-roots and 35% of the container trees. The direction of the basal sweep leaning was concentrated to the NE, which coincided with the direction of the slope and the prevailing wind direction. No effect of planting position in relation to the direction of site preparation was found for basal sweep. The size of the root system, fresh weight and total cross-sectional area was on average for all trees correlated to both DBH and shoot fresh weights. For containers separately, however, only root area was correlated to DBH and stem fresh weight. In spite of the difference in the above-ground size, there were no significant differences in root numbers and root area (CSA) between the two plant types. No difference in rooting depth between plant types was found. When splitting the root system into 120° horizontal sectors (1/3 of the root system) the highest numbers of roots were concentrated in the NW direction. The highest amount of root area tended to be concentrated along a NW-SE diagonal, with a dominance for the latter (SE). Sectors without dominant roots (expected to be the future main structural, stabilizing roots), varied in size (94–178°) but were on average wider in bare-roots and on average concentrated in the NE direction and the downhill side of the slope. This revised version was published online in June 2006 with corrections to the Cover Date.     

Historical biogeography of the ancient lycophyte genus Selaginella: early adaptation to xeric habitats on Pangea

The ancient and cosmopolitan lycophyte genus Selaginella has living representatives around the world, but their historical biogeography has not been assessed with modern methods. We estimated a time‐calibrated phylogeny using DNA marker regions rbcL and ITS1‐5.8S‐ITS2 from 200 species. Node density analyses revealed that Selaginellaceae has significantly older median and mean node ages than other putative “ancient” families. We used statistical model comparison to assess different biogeographical models on our dated tree, and to estimate ancestral ranges. These revealed that Selaginella originated on Euramerica around 383 Ma in the Devonian period, while its peak diversification began with the formation of Pangea. The divergence of the two main species‐rich Selaginella lineages occurred approximately 318 Ma on the supercontinent. The major divergences within these main lineages of Selaginella took place in the Late Permian and Early Triassic, along with lineages highly adapted for xeric habitats on Pangea.     

Root system architecture and gravity perception of a mangrove plant,Sonneratia alba J. Smith

We describe the features of the root system and the gravitropism of roots produced bySonneratia alba. The root system consists of four root types with different growth directions: (a) Pneumatophores, which are negatively orthogravitropic and their statocytes are very large (922 μm²) and the statolith is located near the proximal wall, (b) Cable roots and (c) Feeding roots which are both diagravitropic and their statoliths are settled along the longitudinal wall, and (d) Anchor roots which are positively orthogravitropic. The statocyte cells are the smallest (420 μm²) and statoliths settled at the distal wall. We found that all roots with marked gravitropism have statoliths that settle along different walls of the statocyte. This implies that the statoliths sensing of gravity is done by gravity on mass, and that they are denser than surrounding cytoplasm and this position is related to root growth direction. This finding matches the statoliths sediment under the effect of gravity. Irrespective of statolith, position and direction of growth may be stable.     

Rhacophyton from the Upper Devonian of West Virginia

A new species of Rhacophyton, R. ceratangium , is described from Upper Devonian rocks of West Virginia, U.S.A.; this is synonymous with R. incertum (Dawson) Krausel and Weyland but reasons are cited to indicate that the latter species name is not appropriate.
The collections include: stems up to 2 cm in diameter bearing bipinnate, non-laminate vegetative fronds; abundant well preserved fertile fronds that show clearly the distinctive morphology of their sterile and fertile pinnae; fragments of axes with woody tissues petrified.
The sporangia are particularly distinctive with their long slender tip; they dehisced longitudinally and contained several hundred spores; all available evidence indicates that the plant was homosporous. All petrified axes have a slender bar-shaped strand of primary wood swollen at either end and surrounded by strongly developed secondary wood consisting of scalariform tracheids and rays.
R. ceratangium is closely related to the Belgian R. zygopteroides Leelercq. A comparison with other Devonian and Carboniferous pteridophytes suggests that Rhacophyton is probably a primitive member of the Progymnospermopsida or immediately ancestral to that group.     

The stomatal apparatus of Lycopodium japonicum and its bearing on the stomata of the Devonian lycophyte Drepanophycus spinaeformis

The structure of the stomatal apparatus of the leaf of Lycopodium japonicum Thumb was studied using epidermal macerations, sections and scanning electron microscopy. The stomatal apparatus of L. japonicum consists of two large guard cells and pore, and is anomocytic. Based on light microscopy, the impression from epidermal macerations that there were two small guard cells surrounded by two, large, similarly shaped, subsidiary cells (paracytic) derives from a pronounced elliptical cuticular ledge on the surface of the guard cells surrounding a thickened circumporal area. A similar appearance is characteristic of cuticle preparations of the Devonian lycophyte Drepanophycus spinaeformis Göppert. We therefore conclude, as did W.H. Lang over 70 years ago, that the stomata of the early lycophyte were also anomocytic, as were those of a second species of Drepanophycus , D. qujingensis Li & Edwards.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 209–216.     

Theoretical evidence that root penetration ability interacts with soil compaction regimes to affect nitrate capture

Background and AimsAlthough root penetration of strong soils has been intensively studied at the scale of individual root axes, interactions between soil physical properties and soil foraging by whole plants are less clear. Here we investigate how variation in the penetration ability of distinct root classes and bulk density profiles common to real-world soils interact to affect soil foraging strategies.MethodsWe utilize the functional–structural plant model ‘OpenSimRoot’ to simulate the growth of maize (Zea mays) root systems with variable penetration ability of axial and lateral roots in soils with (1) uniform bulk density, (2) plow pans and (3) increasing bulk density with depth. We also modify the availability and leaching of nitrate to uncover reciprocal interactions between these factors and the capture of mobile resources.Key ResultsSoils with plow pans and bulk density gradients affected overall size, distribution and carbon costs of the root system. Soils with high bulk density at depth impeded rooting depth and reduced leaching of nitrate, thereby improving the coincidence of nitrogen and root length. While increasing penetration ability of either axial or lateral root classes produced root systems of comparable net length, improved penetration of axial roots increased allocation of root length in deeper soil, thereby amplifying N acquisition and shoot biomass. Although enhanced penetration ability of both root classes was associated with greater root system carbon costs, the benefit to plant fitness from improved soil exploration and resource capture offset these.ConclusionsWhile lateral roots comprise the bulk of root length, axial roots function as a scaffold determining the distribution of these laterals. In soils with high soil strength and leaching, root systems with enhanced penetration ability of axial roots have greater distribution of root length at depth, thereby improving capture of mobile resources.     

Enigmatic occurrence of Permian plant roots in Lower Silurian rocks,Guizhou Province,China

Pinnatiramosus qianensis Geng, 1986, is a plant with a complex, extensive pinnate branching system and pitted tracheids, collected from marine Lower Silurian (Llandovery; c. 430 Ma) rocks in Guizhou Province, China. It challenges long‐held theories on the origin and early evolution of vascular plants in the Silurian and Devonian. However, there is a hypothesis that the fossils were not syngenetic with the entombing rock, but were the rooting systems of much younger plants, probably of Permian age. New sections and collections of P. qianensis have been subjected to detailed analyses, which indicate that P. qianensis belongs to an early Permian (c. 285 Ma) rooting system growing down into lower Silurian rocks.     

Fern and lycophyte guard cells do not respond to endogenous abscisic acid

Stomatal guard cells regulate plant photosynthesis and transpiration. Central to the control of seed plant stomatal movement is the phytohormone abscisic acid (ABA); however, differences in the sensitivity of guard cells to this ubiquitous chemical have been reported across land plant lineages. Using a phylogenetic approach to investigate guard cell control, we examined the diversity of stomatal responses to endogenous ABA and leaf water potential during water stress. We show that although all species respond similarly to leaf water deficit in terms of enhanced levels of ABA and closed stomata, the function of fern and lycophyte stomata diverged strongly from seed plant species upon rehydration. When instantaneously rehydrated from a water-stressed state, fern and lycophyte stomata rapidly reopened to predrought levels despite the high levels of endogenous ABA in the leaf. In seed plants under the same conditions, high levels of ABA in the leaf prevented rapid reopening of stomata. We conclude that endogenous ABA synthesized by ferns and lycophytes plays little role in the regulation of transpiration, with stomata passively responsive to leaf water potential. These results support a gradualistic model of stomatal control evolution, offering opportunities for molecular and guard cell biochemical studies to gain further insights into stomatal control.     

A dynamic root system growth model based on L-Systems

Understanding the impact of roots and rhizosphere traits on plant resource efficiency is important, in particular in the light of upcoming shortages of mineral fertilizers and climate change with increasing frequency of droughts. We developed a modular approach to root growth and architecture modelling with a special focus on soil root interactions. The dynamic three-dimensional model is based on L-Systems, rewriting systems well-known in plant architecture modelling. We implemented the model in Matlab in a way that simplifies introducing new features as required. Different kinds of tropisms were implemented as stochastic processes that determine the position of the different roots in space. A simulation study was presented for phosphate uptake by a maize root system in a pot experiment. Different sink terms were derived from the root architecture, and the effects of gravitropism and chemotropism were demonstrated. This root system model is an open and flexible tool which can easily be coupled to different kinds of soil models.     

Strigolactones suppress adventitious rooting in Arabidopsis and pea

Adventitious root formation is essential for the propagation of many commercially important plant species and involves the formation of roots from nonroot tissues such as stems or leaves. Here, we demonstrate that the plant hormone strigolactone suppresses adventitious root formation in Arabidopsis (Arabidopsis thaliana) and pea (Pisum sativum). Strigolactone-deficient and response mutants of both species have enhanced adventitious rooting. CYCLIN B1 expression, an early marker for the initiation of adventitious root primordia in Arabidopsis, is enhanced in more axillary growth2 (max2), a strigolactone response mutant, suggesting that strigolactones restrain the number of adventitious roots by inhibiting the very first formative divisions of the founder cells. Strigolactones and cytokinins appear to act independently to suppress adventitious rooting, as cytokinin mutants are strigolactone responsive and strigolactone mutants are cytokinin responsive. In contrast, the interaction between the strigolactone and auxin signaling pathways in regulating adventitious rooting appears to be more complex. Strigolactone can at least partially revert the stimulatory effect of auxin on adventitious rooting, and auxin can further increase the number of adventitious roots in max mutants. We present a model depicting the interaction of strigolactones, cytokinins, and auxin in regulating adventitious root formation.     

Endophytic cyanobacteria in a 400-million-yr-old land plant: A scenario for the origin of a symbiosis?

Direct evidence for the origin and evolution of land plant/cyanobacterial symbioses is virtually absent from the fossil record. Here we report on rare occurrences of prostrate mycorrhizal axes of the Early Devonian land plant Aglaophyton major that host a filamentous cyanobacterium, which enters the plant through the stomata and colonizes the substomatal chambers and intercellular spaces in the outer cortex. In dead ends of the intercellular system, the filaments form loops and continue growth in reverse direction. Some filaments penetrate parenchyma cells close to and within the mycorrhizal arbuscule-zone and form intracellular coils. This discovery represents the earliest direct evidence for cyanobacteria growing inside land plants, and offers a model for the types of associations that may have preceded the evolution of mutualistic land plant/cyanobacterial symbioses.