Origin of Intra-annual Density Fluctuations in a Semi-arid Area of Northwestern China

Intra-annual density fluctuation (IADF) is a structural modification of the tree ring in response to fluctuations in the weather. The expected changes in monsoon flow would lead to heterogeneous moisture conditions during the growing season and increase the occurrence of IADF in trees of the arid ecosystems of continental Asia. To reveal the timings and physiological mechanisms behind IADF formation, we monitored cambial activity and wood formation in Chinese pine (Pinus tabuliformis) during 2017–2019 at three sites in semi-arid China. We compared the dynamics of xylem formation under a drought event, testing the hypothesis that drought affects the process of cell enlargement and thus induces the production of IADF. Wood microcores collected weekly from April to October were used for anatomical analyses to estimate the timings of cambial activity, and the phases of enlargement, wall thickening, and lignification of the xylem. The first cells started enlargement from late April to early May. The last latewood cells completed differentiation in mid-September. Trees produced IADF in 2018. During that year, a drought in June limited cell production in the cambium, only 36% of the xylem cells being formed in IADF trees, compared to 68% in normal tree rings. IADF cells enlarged under drought in early July and started wall thickening during the rainfall events of late July. The drought restricted cell enlargement and affected wall thickening, resulting in narrow cells with wide walls. Cambium and cell enlargement recovered from the abundant rainfall, producing a new layer with large earlywood tracheids. IADF is a specific adaptation of trees to cope with water deficit events occurring during xylem formation. Our findings confirmed the hypothesis that the June-July drought induces latewood-like IADFs by limiting the process of cell enlargement in the xylem. Our finding suggests a higher occurrence of IADF in trees of arid and semi-arid climates of continental Asia if the changes to monsoon flows result in more frequent drought events during the earlywood formation in June.

Intra-Annual Wood Formation of Cryptomeria fortunei and Cunninghamia lanceolata in Humid Subtropical China

Monitoring cambial activity is important for a better understanding of the mechanisms governing xylem growth responses to climate change, providing a scientific basis for tree-ring-based climate reconstructions and projections about tree growth under future climate scenarios. It plays an even more important role in investigating evergreen tree growth in regions with less distinct seasonal cycles. Subtropical evergreen forests have been studied in recent years for their sensitivity to climate change, but it remains unclear how xylem growth is driven by subtropical climates. To further understand the climate-growth response strategies of subtropical conifers, we micro-cored Cryptomeria fortunei and Cunninghamia lanceolata weekly in 2016 and 2017 at the humid subtropical Gushan Mountain in southeastern China. Our weekly growth monitoring showed that the vegetation periods of these two species were both approximately 2–3 months longer than trees in temperate and boreal forests. The growth of C. fortunei in 2016 and 2017 and C. lanceolata in 2017 showed a bimodal pattern of xylogenesis, which was induced by summer drought. The results also indicated that the earlier end of the xylem formation was related to the yearly drought stress. These findings provide more specific information about tree growth and evidence of how climate influences wood production at the cellular level in subtropical regions.

Description of Intra-Annual Changes in Cambial Activity and Differentiation of Secondary Conductive Tissues of Aesculus hippocastanum Trees Affected by the Leaf Miner Cameraria ohridella

Aesculus hippocastanum trees are commonly infested by the leaf miner Cameraria ohridella, whose larval activity causes the destruction of the leaf parenchyma and induces defoliation. Pest attacks result in, e.g., production of smaller fruits and tree re-flowering in autumn. Concerning pest influence on stem structure only scarce information of narrower annual growth rings of wood has been published. Therefore, we determined the effect of the presence of the leaf miner infestation on intra-annual cambial activity and on differentiation of conductive tissues. These data were compared with phenological phases and pest activity. Pest feeding resulted in changes in onset, cessation and duration of cambial divisions, and differentiation of secondary xylem. The duration of cambial activity was about a month shorter in heavily infested trees and was connected with premature tree defoliation. Affected trees were characterised by a reduction in cambial divisions and earlier cessation of wood differentiation resulting in narrower wood rings. Furthermore, the infested trees exhibited altered wood structure, with more vessels of smaller diameters, however these changes did not affect its theoretical hydraulic conductivity. Interestingly, pest attack did not influence secondary phloem differentiation. The probable influence of long-term infestation on tree growth and condition was discussed.

Cambial Activity of Moringa peregrina (Forssk.) Fiori in Arid Environments

Moringa peregrina (Forssk.) Fiori, one of 13 species of the Moringaceae family widely distributed throughout the dry tropics, has the potential to become one of the most economically important medicinal plants in Egypt. However, despite its tolerance for drought and heat, it is also threatened by increasing temperatures and decreasing precipitation. Although the phenophase of this species is well documented, almost nothing is known about its period of cambial activity in desert regions. Ring formation and the general environmental adaptability of trees are affected by the timing of cambial activation. In our study site, we observe a distinct coupling of the development of new green leaves at the onset of vegetative growth in October and the phase of cambial activity (November–January). The onset of cambial activity seems to be related to a drop in temperature in October and the onset of torrential rains in the region. There might even be a short phase between the end of cambial activity and the onset of bud formation without xylem formation, but with photosynthetic activity. If so, we assume that all assimilates are stored as non-structural carbohydrates (NSC) in the parenchyma of the new ring. This potential gap opens new questions regarding the correlation between NSC storage capacity and the timing of remobilization for subsequent ring formation.

Variation in Onset of Leaf Unfolding and Wood Formation in a Central African Tropical Tree Species

A diversity of phenological strategies has been reported for tropical tree species. Defoliation and seasonal dormancy of cambial activity inform us on how trees cope with water stress during the dry season, or maximize the use of resources during the rainy season. Here, we study the matching between leaf phenology (unfolding and shedding) and cambial activity for Prioria balsamifera, a key timber species in the Democratic Republic of Congo. In particular, we (i) evaluated the seasonality of cambial activity and synchrony of phenology among trees in response to climate and (ii) identified the seasonality of leaf phenology and its relation with cambial phenology. The study was conducted in the Luki Man and Biosphere Reserve, located in the Mayombe forest at the southern margin of the Congo Basin. Historic defoliation data were collected every ten days using weekly crown observations whereas recent observations involved time-lapse cameras. Cambial pinning was performed on ten trees during 20 months and radius dendrometers were installed on three trees during 13 months. Tree rings were measured on cores from 13 trees and growth synchrony was evaluated. We found that P. balsamifera defoliates annually with a peak observed at the end of the dry season and the beginning of the rainy season. The new leaves unfolded shortly after shedding of the old leaves. The peak defoliation dates varied across years from September 12 to November 14 and the fraction of number of trees that defoliated at a given time was found to be negatively correlated with annual rainfall and temperature; during the dry season, when precipitation and temperatures are the lowest. Wood formation (radial growth), was found to be highly seasonal, with cambial dormancy occurring during the dry season and growth starting at the beginning of the rainy season. Individual ring-width series did not cross date well. The within species variability of leaf phenology and cambial rhythms provides indication about resistance of the population against climatic changes.

Transcriptome Analysis of Cambium Tissue of Paulownia Collected during Winter and Spring

Paulownia (Paulownia elongata) is a fast-growing, multipurpose deciduous hardwood species that grows in a wide range of temperatures from –30 °C to 45 °C. Seasonal cues influence the secondary growth of tree stems, including cambial activity, wood chemistry, and transition to latewood formation. In this study, a de novo transcriptome approach was conducted to identify the transcripts expressed in vascular cambial tissue from senescent winter and actively growing spring seasons. An Illumina paired-end sequenced cambial transcriptome generated 297,049,842 clean reads, which finally yielded 61,639 annotated unigenes. Based on non-redundant protein database analyses, Paulownia cambial unigenes shared the highest homology (64.8%) with Erythranthe guttata. KEGG annotation of 35,471 unigenes identified pathways enriched in metabolic activities. Transcriptome-wide DEG analysis showed that 2688 and 7411 genes were upregulated and downregulated, respectively, in spring tissues compared to winter. Interestingly, several transcripts encoding heat shock proteins were upregulated in the spring season. RT-qPCR expression results of fifteen wood-forming candidate genes involved in hemicellulose, cellulose, lignin, auxin, and cytokinin pathways showed that the hemicellulose genes (CSLC4, FUT1, AXY4, GATL1, and IRX19) were significantly upregulated in spring season tissues when compared to winter tissues. In contrast, lignin pathway genes CCR1 and CAD1 were upregulated in winter cambium. Finally, a transcriptome-wide marker analysis identified 11,338 Simple Sequence Repeat (SSRs). The AG/CT dinucleotide repeat predominately represented all SSRs. Altogether, the cambial transcriptomic analysis reported here highlights the molecular events of wood formation during winter and spring. The identification of candidate genes involved in the cambial growth provides a roadmap of wood formation in Paulownia and other trees for the seasonal growth variation.

Triggering Bimodal Radial Stem Growth in Pinus sylvestris at a Drought-Prone Site by Manipulating Stem Carbon Availability

A bimodal radial growth (RG) pattern, i.e., growth peaks in spring and autumn, was repeatedly found in trees in the Mediterranean regions, where summer drought causes reduction or cessation of cambial activity. In a dry inner Alpine valley of the Eastern Alps (Tyrol, Austria, 750 m asl), Pinus sylvestris shows unimodal RG with onset and cessation of cambial activity in early April and late June, respectively. A resumption of cambial activity after intense summer rainfall was not observed in this region. In a field experiment, we tested the hypothesis that early cessation of cambial activity at this drought-prone site is an adaptation to limited water availability leading to an early and irreversible switch of carbon (C) allocation to belowground. To accomplish this, the C status of young P. sylvestris trees was manipulated by physical blockage of phloem transport (girdling) 6 weeks after cessation of cambial cell division. Influence of manipulated C availability on RG was recorded by stem dendrometers, which were mounted above the girdling zone. In response to blockage of phloem flow, resumption of cambial activity was detected above girdling after about 2 weeks. Although the experimentally induced second growth surge lasted for the same period as in spring (c. 2 months), the increment was more than twice as large due to doubling of daily maximum RG rate. After girdling, wood anatomical traits above girdling no longer showed any significant differences between earlywood and latewood tracheids indicating pronounced effects of C availability on cell differentiation. Below girdling, no reactivation of cambial activity occurred, but cell wall thickness of last formed latewood cell was reduced due to lack of C supply after girdling. Intense RG resumption after girdling indicates that cessation of cambial activity can be reversed by manipulating C status of the stem. Hence, our girdling study yielded strong support for the hypothesis that belowground organs exert high C sink strengths on the drought-prone study site. Furthermore, this work highlights the need of in-depth experimental studies in order to understand the interactions between endogenous and exogenous factors on cambial activity and xylem cell differentiation more clearly.

Pharmacognostical and physiochemical evaluation of bruhaniya gana siddha ksheerpak for ksheerpan and ksheerbasti in the management of garbhashosh

Ayurveda is the most ancient medical science framed on evidence-based principles. So, the standardization of Ayurveda drugs is much needed work in view to asses and maintain the quality of Ayurveda drugs in present era. The WHO appreciated the importance of medicinal plants for public health care in developing nations and has evolved guidelines to support the member states in their efforts to formulate national policies on traditional medicine and to study their potential usefulness including evaluation, safety and efficacy. The quality assessment of Ayurveda drug formulations is very important in order to prove their acceptability in modern system of medicine. The present study aims to standardize bruhaniya gana siddha ksheerpaka formulation for ksheerpan and ksheerbasti in the management of garbhashosh (IUGR). The process of standardization is achieved by systematic pharmacognostical and physiochemical evaluative studies. The drug formulation bruhaniya gana siddha ksheer contains Bala, Ashwagandha, Shatawari, Vidari, Kakoli, and Kshirkakoli.2 Organoleptic characters revelled that formulation was light off white in colour, characteristic fragrant odour, and sweet in taste. The result of pharmacognostical study had shown presence of prismatic crystals of Bala, Scalariform vessels of Shatavari, and trichomes of Ashwagandha, cambial activity of the tubers of Vidari, etc. Physiochemical study showed loss on drying 2.96%, total ash 0.31%, acid insoluble ash 0.01%, pH 6.18%, total solids 34.68. Phyto-chemical analysis showed presence of alkaloids, tannins, steroids, proteins, carbohydrates, reducing sugar, etc. The formulation was prepared and screened for various standardization parameters as per ayurveda drug standard protocols in API.

Cambium phenology and dendrochronology of the endangered tropical tree Ocotea catharinensis Mez

Ocotea catharinensis occurs mainly in the Brazilian Atlantic Forest and is highly threatened by timber logging, forest fragmentation, and habitat loss. The remnant populations of this species are genetically rich, which may be related to the presence of long-lived trees, and so it is imperative to understand their growth and age. In this study, we analyzed trees of O. catharinensis from dense ombrophylous forest, its dendrochronological potential, the period of cambial activity and dormancy, and the influence of climate on annual growth. The species showed distinct annually-formed tree-rings, with cambial activity during summer and autumn, and cambial dormancy during winter and spring. A tree-ring width chronology was built from 1852–2015. The estimated age of the sampled trees varied from 40 to 164 years. O. catharinensis in our study site does not show a strong age–diameter relationship, hence similar diameters at breast height resulted in differences in ages by as much as 50 years. The resulting tree-ring width chronology is positively correlated with March precipitation, the transition month between rainy and dry seasons. This study brings valuable contributions to the understanding of the growth of O. catharinensis, which is a novelty for this species and important to the maintenance of these long-lived trees in natural forests.

Cambial Activity and Phenology in Three Understory Species along an Altitude Gradient in Mexico

The aims of this study were to evaluate the cambial activity and phenology of three species with different life forms (Alchemilla procumbens, Acaena elongata and Ribes ciliatum) along an altitudinal gradient and to establish which environmental variables (light, soil humidity and temperature) had the greatest influence on cambial activity and phenological stages. Over two years, data on phenology, growth and cambium were gathered every four weeks in three to six sites per species in Sierra Nevada, Mexico. The results showed that Ribes is the only species that terminates cambial activity with leaves senescence and is influenced by the minimum soil temperature. The light environment influenced the vegetative stages in Alchemilla (cryptophyte), while in Acaena (hemicryptophyte), the mean soil temperature explained the findings related to leaf area during the dry season and growth along the gradient. In the three species, the reproductive phase dominated for a longer period at higher elevations, especially in Alchemilla. Only Ribes, the phanerophyte, showed a similar cambial activity to other trees and shrubs. Although cambium reactivates during the dry season, no xylogenesis occurs. The three species varied during the time in which vascular cambium was active, and this was dependent on the altitude. Specifically, the variation was more rhythmic in Ribes and switched on and off in Alchemilla. It is likely that, depending on the life form, vascular cambium may be more or less susceptible to one or more climate factors.