MIGRATING CAMBIAL DOMAINS AND THE ORIGIN OF WAVY GRAIN IN XYLEM OF BROADLEAVED TREES

Anatomical analysis of wavy-grained xylem samples of Fraxinus, Acer, and Betula reveals a strong underlying pattern of orientational domains within the cambium. Domains are local areas in the cambium within which the orientation of pseudotransverse divisions of fusiform initial cells is predominately unidirectional rather than random. The orientation of ray splitting and ray uniting also tends to be unidirectional within domains. Because their boundaries migrate with time, domains are not fixed in shape or position. A specific group of initial cells may first be located in the interior of a domain, later near a boundary, and still later in another domain of opposite orientational prevalence. Domain patterns and their movements were delineated by classifying and mapping ray splitting and uniting events in temporal series of tangential sections of terminal late xylem. The domain patterns move slowly upward, preceding the similarly outlined, but noncoincident, grain pattern. A wavy grain pattern can be interpreted as being the result of a domain pattern of similar geometry. The efficacy of a domain pattern in eliciting a grain pattern is probably indirectly dependent upon the frequency of occurrence of pseudotransverse divisions and of ray splitting and uniting. The frequency is low in cambium producing straight-grained wood. Evidence of 10-fold and greater increases in frequency of ray splitting and uniting was found in straight-grained to wavy-grained transitional xylem.     

SUCCESSIVE CAMBIA IN THE STEM OF PHYTOLACCA DIOICA

Phytolacca dioica L., an evergreen tree of the Phytolaccaceae, is one of the species of Phytolacca which shows anomalous secondary thickening in its stem. This mode of thickening has been regarded as successive cambial activity or alternatively, in some more recent interpretations, as thickening by unidirectional activity of a cambial zone. The stem thickening of P. dioica is of the former type. The cambium produces fascicular strands, showing centrifugal differentiation of xylem and centripetal differentiation of phloem on opposite sides of the cambial layer, and rays are produced between the fascicular areas. In both xylem and phloem the younger elements are closer to the cambium than the older elements. Succeeding cambia arise periodically by periclinal divisions in a layer of parenchyma cells two or three cells beyond the outermost intact phloem derived from the current cambium. Each cambium forms a few parenchyma cells on both sides before it forms derivatives which mature into lignified xylem elements or conductive elements of the phloem. The parenchyma thus formed toward the outside later becomes the site of the origin of the succeeding cambium. Only one or two layers of this phloem parenchyma go on to form the new cambium; the remaining cells accumulate between the outermost phloem and the cortex. P. weberbaueri shows stem structure similar to P. dioica. P. meziana, a shrub, shows normal stem structure.     

Amphibian survival,growth and development in response to mineral nitrogen exposure and predator cues in the field: an experimental approach

Mineral nitrogen (N) has been suggested as a potential factor causing declines in amphibian populations, especially in agricultural landscapes; however, there is a question as to whether it remains in the water column long enough to be toxic. We explored the hypothesis that mineral N can cause both lethal and sublethal toxic effects in amphibian embryos and larvae in a manipulative field experiment. We sampled 12 ponds, fertilizing half with ammonium nitrate fertilizer early in the spring, and measured hatching, survival, development, growth, and the incidence of deformities in native populations of wood frog (Rana sylvatica) and eastern tiger salamander (Ambystoma tigrinum tigrinum) embryos and larvae held in in situ enclosures. We found that higher ammonium concentrations negatively affect R. sylvatica more strongly than A. tigrinum. R. sylvatica tended to have lower survival as embryos and young tadpoles, slowed embryonic development, and an increased proportion of hatchlings with deformities at experimentally elevated ammonium. A. tigrinum did not experience significantly reduced survival, but their larval development was slowed in response to elevated ammonium and the abundance of large invertebrate predators. Variable species susceptibility, such as that shown by R sylvatica and A. tigrinum, could have large indirect effects on aquatic community structure through modification of competitive or predator-prey relationships. Ammonium and nitrate + nitrite concentrations were not correlated with other measures that might have affected amphibians, such as pH, pond area, depth, or vegetation. Our results highlight the potential importance of elevated ammonium on the growth, development and survival of amphibians, especially those that breed in surface waters receiving anthropogenic N inputs. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

MOL1 is required for cambium homeostasis in Arabidopsis

Plants maintain pools of pluripotent stem cells which allow them to constantly produce new tissues and organs. Stem cell homeostasis in shoot and root tips depends on negative regulation by ligand–receptor pairs of the CLE peptide and leucine‐rich repeat receptor‐like kinase (LRR‐RLK) families. However, regulation of the cambium, the stem cell niche required for lateral growth of shoots and roots, is poorly characterized. Here we show that the LRR‐RLK MOL1 is necessary for cambium homeostasis in Arabidopsis thaliana. By employing promoter reporter lines, we reveal that MOL1 is active in a domain that is distinct from the domain of the positively acting CLE41/PXY signaling module. In particular, we show that MOL1 acts in an opposing manner to the CLE41/PXY module and that changing the domain or level of MOL1 expression both result in disturbed cambium organization. Underlining discrete roles of MOL1 and PXY, both LRR‐RLKs are not able to replace each other when their expression domains are interchanged. Furthermore, MOL1 but not PXY is able to rescue CLV1 deficiency in the shoot apical meristem. By identifying genes mis‐expressed in mol1 mutants, we demonstrate that MOL1 represses genes associated with stress‐related ethylene and jasmonic acid hormone signaling pathways which have known roles in coordinating lateral growth of the Arabidopsis stem. Our findings provide evidence that common regulatory mechanisms in different plant stem cell niches are adapted to specific niche anatomies and emphasize the importance of a complex spatial organization of intercellular signaling cascades for a strictly bidirectional tissue production.     

Carboniferous Onychophora from Montceau‐les‐Mines,France, and onychophoran terrestrialization

The geological age of the onychophoran crown‐group, and when the group came onto land, have been sources of debate. Although stem‐group Onychophora have been identified from as early as the Cambrian, the sparse record of terrestrial taxa from before the Cretaceous is subject to contradictory interpretations. A Late Carboniferous species from the Mazon Creek biota of the USA, Helenodora inopinata, originally interpreted as a crown‐group onychophoran, has recently been allied to early Cambrian stem‐group taxa. Here we describe a fossil species from the Late Carboniferous Montceau‐les‐Mines Lagerstätte, France, informally referred to as an onychophoran for more than 30 years. The onychophoran affinities of Antennipatus montceauensis gen. nov., sp. nov. are indicated by the form of the trunk plicae and the shape and spacing of their papillae, details of antennal annuli, and the presence of putative slime papillae. The poor preservation of several key systematic characters for extant Onychophora, however, prohibits the precise placement of the Carboniferous fossil in the stem or crown of the two extant families, or the onychophoran stem‐group as a whole. Nevertheless, A. montceauensis is the most compelling candidate to date for a terrestrial Paleozoic onychophoran.     

Growth and posture control strategies in Fagus sylvatica and Acer pseudoplatanus saplings in response to canopy disturbance

Background and Aims

Forest tree saplings that grow in the understorey undergo frequent changes in their light environment to which they must adapt to ensure their survival and growth. Crown architecture, which plays a critical role in light capture and mechanical stability, is a major component of sapling adaptation to canopy disturbance. Shade-adapted saplings typically have plagiotropic stems and branches. After canopy opening, they need to develop more erect shoots in order to exploit the new light conditions. The objective of this study was to test whether changes in sapling stem inclination occur after canopy opening, and to analyse the morphological changes associated with stem reorientation.

Methods

A 4-year canopy-opening field experiment with naturally regenerated Fagus sylvatica and Acer pseudoplatanus saplings was conducted. The appearance of new stem axes, stem basal diameter and inclination along the stem were recorded every year after canopy opening.

Key Results

Both species showed considerable stem reorientation resulting primarily from uprighting (more erect) shoot movements in Fagus, and from uprighting movements, shoot elongation and formation of relay shoots in Acer. In both species, the magnitude of shoot uprighting movements was primarily related to initial stem inclination. Both the basal part and the apical part of the stem contributed to uprighting movements. Stem movements did not appear to be limited by stem size or by stem growth.

Conclusions

Stem uprighting movements in shade-adapted Fagus and Acer saplings following canopy disturbance were considerable and rapid, suggesting that stem reorientation processes play a significant role in the growth strategy of the species.     

L'evoluzione della cerchia legnosa in Arbutus Andrachnoides Link. in Firenze,dal luglio 1946 al Giugno 1947

Summary

The evolution of the wood ring in arbutus andrachnoides link, in florence (from July 1946 to June 1947).

The anatomical characters of the wood ring of the stem and of the young branch of Arbutus andrachnoides Link have been studied in Florence.

The results are the following: the cambial tissue in the stem starts dividing at the end of April and continues during the summer months, ending completely in September.

In the branch the cambium differentiates almost a month earlier, namely at the beginning of April.

In both cases the only wood ring produced is completed at the end of September. Comparing the evolution of this wood with the climatic factors it can be seen that the cambium activity starts in a period of remarkable rainfall and of regular increase of temperature and presents a remarkable activity still at the end of August.

Arbutus andrachnoides Link. is a spontaneous hybrid between A. Unedo and A. Andrachne and its distribution (in Greece and in the gréek islands) shows that A. Unedo and A. Andrachne had the possibility to overlap in this area and to produce this hybrid.     

Andamento Dell'Accrescimento Del Legno in Ceratonia Siliqua L.

Abstract

The woody tissue of Ceratonia siliqua is of the diffuse-porous type, being mainly constituted of fibres full of starch grains (sostitution fibres; fusiform cells of the wood parenchyma).

The wood formed throughout the year is uniform and the annual rings are hardly distinguishable except for the presence of apotracheal parenchyma. The cambium shows a marked activity from May to July, and a less pronounced one from July to September; it is hardly or not at all active from October to April. During these latter months the cambium is completely dormant only in January, while during the others, especially the ones prior to the late spring reawakening, it can function in localised areas for short lapses of time.

These observations have been carried out in a female plant cultivated in Apulia, a region where Ceratonia siliqua is not a normal component of the natural plant cover. It is of some interest therefore to compare them with what is known of the same species in Israel, where C. siliqua is a spontaneous element of the maquis vegetation. The behaviour of the cambium of C. siliqua is similar in both regions but in Israel the maximum of activity begins earlier in the year and last for a longer period (mid-April to mid- August), after which the cambium becomes gradually less active until its activity is reduced to a minimum during autumn and winter. A short period of dormancy precedes the spring-time reawakening. This subcontinuous activity of the cambium together with the occurrence of flowering time in autumn-winter shows C. siliqua to be a tropical type of plant, but is nevertheless in good agreement with the eastern mediterranean climate.

In Apulia on the other hand we notice that the subcontinuous activity of the cambium is more potential than real, giving rise to sporadic and localised episodes from October to May. While this and other differences can be interpreted as direct responses to local climatic conditions, the general disturbances in the vegetative cycle can not be explained in the same way. In other words while the tendency of the cambium to remain dormant from October to May is probably caused by the relatively severe winter conditions, the shorter period of functioning in the spring—summer, the full resting period occurring in January rather than in February-March, and the attempts to resume meristematic activity already in February, while the full reawakening only occurs in May, can only be ascribed to indirect climatic action. These disturbances are probably to be seen as conseguences of the fact that the climatic conditions during the Apulian summer allow the flowers, whose primordia are well evident in July, to blossom in October-November and exceptionally even in August that is at least one month earlier than in Israel, where flowering occurs in December. This alteration in the flowering time affects the rythm of cambial activity causing it to be out of phase with the climatic cycle.

C. siliqua appears therefore not to be perfectly adapted to the Apulian climate even though it can often be found where it could not possibly have been planted, as on the steep flanks of fairly deep gulleys, and can be considered to be at least sub-spontaneous.     

Fertility tracers and monsoon forcing at an equatorial site of the Somali Basin (Northwest Indian Ocean)

Temporal changes in biological and isotopic tracers have successfully been used to monitor monsoon variability in the Arabian Sea. Convincing evidence that fertility indicators, such as the relative abundance of the planktic foraminifer Globigerina bulloides, and SiO₂ and Ba fluxes, can monitor monsoon changes, has also been published. Time series of the ¹³C content of the thermocline foraminifer Neogloboquadrina dutertrei, and of an Upwelling Radiolarian Index (URI), have been used to reconstruct upwelling changes in the Somali Basin at 5 °N for the last 160 kyr. In order to establish a reference site for comparison with other upwelling and/or fertility records from the same area, a time series of 8 climatic and fertility proxies (¹³C content of N. dutertrei and Globorotalia menardii, percentages of G. bulloides and of a thermocline foraminiferal group, minimal δ¹⁸ values of Globigerinoides sacculifer, N. dutertrei and G. menardii) are investigated in the spectral domain for the last 360 kyr, at a site located at the equator and outside the Somali upwelling centers (Core MD 85668).Chronostratigraphies for these records are developed by correlation of oxygen isotope record of G. sacculifer to the Martinson standard stacked benthic oxygen isotope record. The temporal resolution of the data, is between 2 and 5 kyr.Cross spectral comparisons with ETP (Eccentricity + Tilt + Precession-composite signal) were used to estimate coherences in conjunction with phase relationships and to quantify relationships between fertility and climatic indicators. The timing of fertility indicator cycles at the equator shows little (obliquity band) or no (other orbital bands) coincidence with monsoon proxies of the Arabian Sea. At the equator, fertility responses are dissociated, and present important leads or lags with ETP signal at the three orbital bands (with the exception of the thermocline foraminifera, in phase with maximum eccentricity). Due to significant leads with ETP in the timing of ¹³C minima and URI cycles, no relationships with the Northern Hemisphere radiation can be deduced. At the obliquity and precessional frequency bands, thermocline foraminifera cycles display significant lag with ETP and ice volume minima, which suggests that radiation is not the sole forcing mechanism. Global climatic and ice volume changes may be an important forcing mechanism for these fertility proxies. The G. bulloides record is more complicated with cycles in phase with Arabian Sea monsoon proxies at the obliquity band, and ¹⁸O minima at the precession frequency.It appears that, at the equator, the southwest monsoon winds are not the main force driving radiolarian and foraminiferal productivity.     

Auxin Stimulation of Cambial Activity in Pinus silvestris I. The Differential Cambial Response

Isolated stem segments of Pinus silvestris L. produce new xylem in sterile culture for 5 weeks if sucrose and IAA are present in the medium. The response of cambium varies in the course of the season and along the tree stem. The cambium is more sensitive in spring and in the stem portion closer to tree apex than later in the season and closer to the stem base. Spring initiation of cambial activity in adult pine trees under natural conditions could not be correlated with any consistent concentration gradient of natural auxin extracted from the cambial region. Thus, the relation between concentration of auxin and the activity of cambium is complex and involves changes of cambial responsivity. Interaction with gibberellic acid or kinetin and changing concentration of sucrose were studied during the season, but none of these substances alone appeared to be responsible for the observed variation in cambial response to auxin.     

Density of an intraguild predator mediates feeding group size, intraguild egg predation, and intra- and interspecific competition

Intraguild predation (IGP) is common in most communities, but many aspects of density-dependent interactions of IG predators with IG prey are poorly resolved. Here, we examine how the density of an IG predator can affect feeding group size, IG egg predation, and the growth responses of IG prey. We used laboratory feeding trials and outdoor mesocosm experiments to study interactions between a social intraguild predator (larvae of the wood frog; Rana sylvatica) and its prey (spotted salamander; Ambystoma maculatum). Larvae of R. sylvatica could potentially affect A. maculatum by consuming shared larval food resources or by consuming eggs and hatchlings. However, successful egg predation requires group feeding by schooling tadpoles. We established from five to 1,190 hatchlings of R. sylvatica in mesocosms, then added either 20 A. maculatum hatchlings to study interspecific competition, or a single egg mass to examine IGP. Crowding strongly suppressed the growth of R. sylvatica, and IGP was restricted to the egg stage. In the larval competition experiment, growth of A. maculatum was inversely proportional to R. sylvatica density. In the predation experiment, embryonic mortality of A. maculatum was directly proportional to the initial density of R. sylvatica and the mean number of tadpoles foraging on egg masses. IGP on eggs reduced A. maculatum hatchling density, which accelerated larval growth. Surprisingly, the density of R. sylvatica had no overall effect on A. maculatum growth because release from intraspecific competition via egg predation was balanced by increased interspecific competition. Our results demonstrate that the density of a social IG predator can strongly influence the nature and intensity of interactions with a second guild member by simultaneously altering the intensity of IGP and intra- and interspecific competition.L . A. Burley and A. T. Moyer contributed equally to this work.     

Targeting Ascophyllum nodosum (L.) Le Jol. extract application at five growth stages of winter wheat

Seaweed extracts are a category of biostimulant products that have the potential to promote plant growth and thus, have given a promise to develop more resilient systems of crop production that make efficient use of water and nutrients. This field study evaluated the effect of application timing of an Ascophyllum nodosum extract on nutrient uptake and grain yield of winter wheat when the extract was applied singly in five growth stages from 2017 to 2019. Analysis by orthogonal polynomial contrasts indicated that extract application significantly increased yield or biomass in all three site-years. Application at the tillering stage increased average yield by 17%, grain nutrient accumulation (N, P, K) by 21–24%, and N-use efficiency by 11% over the three site-year period, but extract effects were not consistent between site-years both in terms of optimal growth stages of application and magnitude of crop responses. While grain yield increased by 39% with extract application at tillering in 2017, yield gains ranging from 16 to 22% were obtained at the stem elongation and booting stages in 2018. These differences were likely the result of site-year variations in weather patterns and soil properties. Among macro-nutrients, only grain N accumulation was linearly correlated to grain yield across site-years indicating that increased soil N uptake and/or remobilization to the reproductive organs was a key process of the A. nodosum mode of action. An economic analysis indicated that the average return over extract cost was higher than that of the zero-extract control by €79-100 ha^-1 when the extract was applied at the tillering or stem elongation stage and assuming higher dilution rates recommended by the industry. Even greater economic benefits can result if N rates are substantially reduced without an adverse effect on yield gains.

     

The genetics of nitrogen use in hexaploid wheat: N utilisation, development and yield

A genetic study is presented for traits relating to nitrogen use in wheat. Quantitative trait loci (QTLs) were established for 21 traits relating to growth, yield and leaf nitrogen (N) assimilation during grain fill in hexaploid wheat (Triticum aestivum L.) using a mapping population from the cross Chinese Spring × SQ1. Glutamine synthetase (GS) isozymes and estimated locations of 126 genes were placed on the genetic map. QTLs for flag leaf GS activity, soluble protein, extract colour and fresh weight were found in similar regions implying shared control of leaf metabolism and leaf size. Flag leaf traits were negatively associated with days to anthesis both phenotypically and genetically, demonstrating the complex interactions of metabolism with development. One QTL cluster for GS activity co-localised with a GS2 gene mapped on chromosome 2A, and another with the mapped GSr gene on 4A. QTLs for GS activity were invariably co-localised with those for grain N, with increased activity associated with higher grain N, but with no or negative correlations with grain yield components. Peduncle N was positively correlated, and QTLs co-localised, with grain N and flag leaf N assimilatory traits, suggesting that stem N can be indicative of grain N status in wheat. A major QTL for ear number per plant was identified on chromosome 6B which was negatively co-localised with leaf fresh weight, peduncle N, grain N and grain yield. This locus is involved in processes defining the control of tiller number and consequently assimilate partitioning and deserves further examination. Electronic Supplementary Material The online version of this article () contains supplementary material, which is available to authorized users.     

Mechanisms and genetic diversity for host plant resistance to spotted stem borer,Chilo partellus in sorghum,Sorghum bicolor

Levels of resistance to spotted stem borer, Chilo partellus, in the germplasm are low to moderate and therefore we evaluated 25 sorghum genotypes for resistance to stem borer to identify lines with diverse mechanisms of resistance to this insect. Leaf glossiness was significantly and negatively associated with low deadheart formation. Dwarf genotypes with fewer nodes showed more deadheart formation. Antixenosis and/or antibiosis for leaf feeding, and reduced deadheart formation, tunnelling, and number of exit holes resulted in lower losses owing to C. partellus damage. Axillary tillers compensated for loss in grain yield owing to borer infestation as a result of deadheart formation, but their synchrony for maturity with the main plant is quite important. Path coefficient analysis revealed that direct effects of stem tunnelling on loss in grain yield were greater than leaf feeding and deadhearts. However, leaf feeding via stem tunnelling showed maximum indirect effects on loss in grain yield. Estimates for broad‐sense heritability and genetic advance suggested that it is possible to improve for grain yield under stem borer infestation. Heritability estimates for grain yield were usually higher than for stem borer damage parameters. Multi‐trait cluster analysis placed the test genotypes into four and five clusters in rainy and post‐rainy seasons, respectively. The genotypes placed in different groups, and showing resistance to leaf feeding, deadheart formation, stem tunnelling, and/or compensation in grain yield can be used for sorghum improvement.     

EVOLUTIONARY HISTORY OF NORTHERN HEMISPHERE NUCELLA (GASTROPODA,MURICIDAE): MOLECULAR,MORPHOLOGICAL, ECOLOGICAL,AND PALEONTOLOGICAL EVIDENCE

By combining data from a variety of sources we explore patterns of evolution and speciation in Nucella, a widely studied genus of shallow-water marine neogastropods. We present a hypothesis of phylogenetic relationships for all of the currently recognized species of northern hemisphere Nucella, based on an analysis of 718 base pairs of nucleotide sequence from the mitochondrial cytochrome b gene. The order of appearance of species in the fossil record is congruent with this hypothesis. The topology of the inferred phylogeny of Nucella, coupled with ecological, morphological, and fossil evidence, was used to address three main questions: (1) At what time and by which route was the North Atlantic invaded from the North Pacific compared to prior studies of the trans-Arctic interchange? (2) Do patterns of molecular variation within species corroborate the importance of climatic cycles in driving speciation in north temperate marine animals? (3) Was radiation in the direction of increased or decreased ecological specialization, body size, or vulnerability to predation? Molecular evidence confirmed that the sole North Atlantic species, N. lapillus, arose from a North Pacific ancestor. Biogeographic and paleontological evidence supported the dispersal of Nucella, and perhaps other interchange species, via the Eurasian Arctic. Rather intriguingly, the linkage of N. lapillus to a western as opposed to eastern Pacific clade, and the biogeographic origins of the eastern Pacific species, parallel closely similar patterns observed in another genus of rocky-shore gastropods, Littorina. This congruence, in conjunction with information on the climatic and geographic histories of the region, as well as the geographic arrangement of mtDNA haplotypes within Nucella species, supports a model of speciation in Nucella driven by cycles of climatic amelioration and deterioration that began during the Miocene. Calibrations from the fossil record of Nucella suggest that third position transitions and transversions accrue at a rate of 3–4% and 0.5% respectively per million yr. This supports an early participation by Nucella in the trans-Arctic interchange, as suggested by paleobiogeographic studies. Consistent with the unstable taxonomic history of species of Nucella, we found few nonmolecular traits to be phylogenetically informative. Among North Pacific species, more recently derived species (N. canaliculata and the N. emarginata clade) were more ecologically specialized (narrower diet and habitat range). Consistent with extensive intraspecific variation, shell traits were quite labile evolutionarily: neither overall size nor development of antipredatory traits exhibited consistent evolutionary trends over the history of the genus. Nurse eggs (unfertilized eggs consumed by developing embryos) were an ancestral trait that was lost evolutionarily in the two clades that also exhibited increased body size, suggesting that these two life-history traits may be coupled. The reduced number of chromosomes in N. lapillus is clearly a derived state and is consistent with White's (1978) observations on chromosome evolution in other clades.     

The Fagus sylvatica forests in the Larvik region, south-eastern Norway: their origin and history

The origin and developmental history of Fagus sylvatica forests in south-eastern Norway have been studied through pollen analysis and AMS radiocarbon-dating of peat from two small forest hollows. In this area F. sylvatica appears to have a long history, from the first occurrence of F. sylvatica pollen at ca. 9100 cal. b.p. to its local expansion ca. 1300–1200 cal. b.p. At this time a shift from a diverse landscape mosaic with many plant taxa present, including broad-leaved trees, to a less diverse landscape mosaic with Picea abies and F. sylvatica trees is interpreted from the pollen data. The long history of F. sylvatica suggests that the existing forests are not recent plantations, but implies that these forests are native. The presence of pollen indicative of anthropogenic activity combined with charcoal before the expansion of F. sylvatica, as well as comparison with data from nearby sites, suggest that the forest development was likely to be a result of human activity and climatic changes, particularly changes in moisture conditions.