Questions
Steppes are among the most transformed and fragmented biomes in the world, but studies on the impact of habitat loss and fragmentation on these zonal grasslands are scarce. In many regions, ...steppe has survived only on small habitat islands, particularly on kurgans (ancient burial mounds) surrounded by intensive croplands. We asked how the size of kurgans, their isolation and other landscape and local factors influence their species richness.
Location
Southern Ukraine.
Methods
We recorded all species of vascular plants and assessed local (area, steepness, disturbance level), landscape (distance to the nearest habitat patch, habitat amount in the surroundings, surrounding type, distance to settlements) and climatic (precipitation) conditions of 112 kurgans. We analysed species–area relationships (SARs) for total richness, habitat specialists and generalists. Then, we assessed importance of environmental factors in explaining the residuals of the SAR models.
Results
Patch area explained more than 50% of variance in total and specialist species richness. The slopes of the power function SAR (z‐values) were higher for specialists than for generalists. Connectivity measures (the distance to the nearest large habitat enclave and share of habitat in the surroundings) were more important for richness of specialists than for that of generalists, which were more controlled by the kurgan's surroundings.
Conclusions
We found that for specialist species the steppe “islands” follow the theory of island biogeography closely, but less so for generalist species. Combined with the revealed importance of connectivity for specialist species richness, we conclude that the conservation value of the kurgans in the long term likely can only be achieved with larger steppe areas in the surroundings.
Kurgans (ancient burial mounds) constitute small steppe habitat islands within the “sea of fields” forming the landscape of southern Ukraine. We found that they can function nearly as "oceanic islands," particularly for steppe vascular plant species as their richness is strongly controlled and positively related with kurgan size, landscape‐scale connectivity and local‐scale habitat heterogeneity indices.
Civilizations, including ancient ones, have shaped global ecosystems in many ways through coevolution of landscapes and humans. However, the cultural legacies of ancient and lost civilizations are ...rarely considered in the conservation of the Eurasian steppe biome. We used a data set containing more than 1000 records on localities, land cover, protection status, and cultural values related to ancient steppic burial mounds (kurgans); we evaluated how these iconic and widespread landmarks can contribute to grassland conservation in the Eurasian steppes, which is one of the most endangered biomes on Earth. Using Bayesian logistic generalized regressions and proportional odds logistic regressions, we examined the potential of mounds to preserve grasslands in landscapes with different levels of land‐use transformation. We also compared the conservation potential of mounds inside and outside protected areas and assessed whether local cultural values support the maintenance of grasslands on them. Kurgans were of great importance in preserving grasslands in transformed landscapes outside protected areas, where they sometimes acted as habitat islands that contributed to habitat conservation and improved habitat connectivity. In addition to steep slopes hindering ploughing, when mounds had cultural value for local communities, the probability of grassland occurrence on kurgans almost doubled. Because the estimated number of steppic mounds is about 600,000 and similar historical features exist on all continents, our results may be applicable at a global level. Our results also suggested that an integrative socioecological approach in conservation might support the positive synergistic effects of conservation, landscape, and cultural values.
Contribución de los valores culturales para la conservación esteparia en los antiguos montículos funerarios de Eurasia
Resumen
Las civilizaciones modernas y antiguas han moldeado de muchas maneras los ecosistemas globales mediante la coevolución del paisaje y la humanidad. Sin embargo, pocas veces se considera el legado cultural de las civilizaciones perdidas o antiguas para la conservación del bioma de la estepa euroasiática. Usamos un conjunto de datos que contiene más de 1,000 registros de las localidades, cobertura del suelo, estado de protección y valores culturales relacionados con los antiguos montículos funerarios de esta estepa (kurgans). Después analizamos cómo estos símbolos icónicos y distribuidos extensamente pueden contribuir a la conservación de los pastizales en la estepa euroasiática, uno de los biomas en mayor peligro de extinción. Analizamos el potencial de conservación de los montículos en paisajes con diferentes niveles de transformación en el uso de suelo mediante regresiones logísticas generalizadas bayesianas y regresiones logísticas de probabilidades proporcionales. También comparamos el potencial de conservación de los montículos dentro y fuera de las áreas protegidas y evaluamos si los valores culturales locales conservan los pastizales dentro de estas mismas áreas. Los kurgans fueron de gran importancia para la conservación de los pastizales en los paisajes transformados ubicados fuera de las áreas protegidas, en donde llegaron a fungir como hábitats aislados que contribuyeron a la conservación y conectividad del hábitat. Además de que las pendientes pronunciadas impiden el arado, cuando los montículos contaban con valor cultural para las comunidades locales, la probabilidad de que el pastizal se ubicara sobre un kurgan casi se duplicó. Ya que se estima que el número de montículos esteparios ronda los 6,000 y que rasgos históricos similares existen en todos los continentes, nuestros resultados pueden aplicarse a nivel global. Nuestros resultados también sugieren que una estrategia socio‐ecológica integradora para la conservación podría respaldar los efectos sinérgicos positivos de la conservación, el paisaje y los valores culturales.
Questions
European steppes are among the most threatened ecosystems in the Palaearctic region, mainly because of conversion to arable land. Abandonment may allow for the passive recovery of steppes. ...We made use of an exceptional old‐field succession chronosequence of nearly 100 years to answer the following questions: (a) Are the plant species composition, species richness and functional characteristics typical of virgin grass steppes able to self‐restore during ca. 100 years after abandonment? (b) Do the rates of recovery of the above vegetation characteristics differ over the studied chronosequence? (c) Do topsoil carbon and nitrogen content change over the succession chronosequence, leading to concentrations similar to that of virgin steppes?
Location
Southern Ukraine.
Methods
We sampled vegetation and soil in a virgin grass steppe and in old fields abandoned for 6, 15, 31, 50 and ca. 97 years. We subjected the composition data to multivariate analysis. To test whether species richness, functional and soil characteristics of the old fields diverge from those of the virgin steppe, we used one‐way analysis of variance with Tukey's honestly significant difference (HSD) statistic to create 90% confidence intervals.
Results
The vegetation composition of the three most recently abandoned old fields differed significantly from that of the virgin steppe. The species richness of vascular plants was lower in old fields than in the virgin steppe. The share of steppe habitat specialists was similar to the virgin steppe only in the field abandoned for ca. 97 years. Functional characteristics were significantly different from the virgin steppe only in the most recently abandoned old field. Contents of Corg and Ntot in fields abandoned for ≤50 years were lower compared with the virgin steppe.
Conclusions
The functional characteristics of steppe vegetation seem to recover much faster than its biodiversity. However, based on our results, 100 years can be enough time for the spontaneous re‐establishment of typical steppe vegetation.
In southern Ukraine, we sampled vegetation and soil in virgin grass steppe and old fields aged from 6 to ca. 97 years. The functional characteristics of steppe vegetation seem to recover much faster than its biodiversity and soil properties. However, based on our results, 100 years can be enough for such an ecosystem to self‐restore.
Auxin is an important regulator of plant ontogenies including embryo development and the exogenous application of this phytohormone has been found to be necessary for the induction of the embryogenic ...response in plant explants that have been cultured
. However, in the present study, we show that treatment of Arabidopsis explants with trichostatin A (TSA), which is a chemical inhibitor of histone deacetylases, induces somatic embryogenesis (SE) without the exogenous application of auxin. We found that the TSA-treated explants generated somatic embryos that developed efficiently on the adaxial side of the cotyledons, which are the parts of an explant that are involved in auxin-induced SE. A substantial reduction in the activity of histone deacetylase (HDAC) was observed in the TSA-treated explants, thus confirming a histone acetylation-related mechanism of the TSA-promoted embryogenic response. Unexpectedly, the embryogenic effect of TSA was lower on the auxin-supplemented media and this finding further suggests an auxin-related mechanism of TSA-induced SE. Congruently, we found a significantly increased content of indolic compounds, which is indicative of IAA and an enhanced DR5::GUS signal in the TSA-treated explants. In line with these results, two of the
genes (
and
), which are involved in auxin biosynthesis, were found to be distinctly up-regulated during TSA-induced SE and their expression was colocalised with the explant sites that are involved in SE. Beside auxin, ROS were extensively accumulated in response to TSA, thereby indicating that a stress-response is involved in TSA-triggered SE. Relevantly, we showed that the genes encoding the transcription factors (TFs) that have a regulatory function in auxin biosynthesis including
, and stress responses (
) were highly up-regulated in the TSA-treated explants. Collectively, the results provide several pieces of evidence about the similarities between the molecular pathways of SE induction that are triggered by TSA and 2,4-D that involve the activation of the auxin-responsive
genes that have a regulatory function in auxin biosynthesis and stress responses. The study suggests the involvement of histone acetylation in the auxin-mediated release of the embryogenic program of development in the somatic cells of Arabidopsis.
The LEAFY COTYLEDON2 (LEC2) transcription factor with a plant-specific B3 domain plays a central role in zygotic and somatic embryogenesis (SE). LEC2 overexpression induced in planta leads to ...spontaneous somatic embryo formation, but impairs the embryogenie response of expiants cultured in vitro under auxin treatment. The auxin-related functions of LEC2 appear during SE induction, and the aim of the present study was to gain further insights into this phenomenon. To this end, the effect of LEC2 overexpression on the morphogenic responses of Arabidopsis expiants cultured in vitro under different auxin treatments was evaluated. The expression profiles of the auxin biosynthesis genes were analysed in embryogenie cultures with respect to LEC2 activity. The results showed that LEC2 overexpression severely modifies the requirement of cultured expiants for an exogenous auxin concentration at a level that is effective in SE induction and suggested an increase in the auxin content in 35S::LEC2-GR transgenic expiants. The assumption of an LEC2 promoted increase in endogenous auxin in cultured expiants was further supported by the expression profiling of the genes involved in auxin biosynthesis. The analysis indicated that YUCCAs and TAA1, working in the IPAYUC auxin biosynthesis pathway, are associated with SE induction, and that the expression of three YUCCA genes (YUC1, YUC4 and YUC10) is associated with LEC2 activity. The results also suggest that the 1AOx-mediated auxin biosynthesis pathway involving ATR1/MYB34 and CYP79B2 does not seem to be involved in SE induction. We conclude that de novo auxin production via the tryptophan-dependent IPA-YUC auxin biosynthesis pathway is implicated in SE induction, and that LEC2 plays a key role in this mechanism.
Steppe is among the most endangered biomes of the world, especially in Eastern Europe, where more than 90 % of original steppes have been destroyed due to conversion into croplands, afforestation and ...other human activities. Currently, steppe vegetation is often restricted to places unsuitable for ploughing, such as ancient burial mounds called kurgans. The aim of our study was to collect and synthesise existing knowledge on kurgans by a review of research papers and grey literature. The proportion of kurgans covered by steppe vegetation increases from west to east and from lowlands to uplands. Despite their small size, kurgans act as biodiversity hotspots and harbour many red-listed species. High overall species richness and a high proportion of grassland specialists are maintained by a pronounced fine-scale environmental heterogeneity. The main factors threatening the biodiversity of kurgans are intensified agriculture and construction works. We conclude that kurgans can play a crucial role in preserving steppe vegetation, especially in intensively used agricultural landscapes in the western part of the steppe zone. Despite the vital role of kurgans in sustaining steppe vegetation, we identified serious knowledge gaps on their distribution, vegetation, flora and fauna and their potential role in steppe restoration.
The angiosperm seed is composed of three genetically distinct tissues: the diploid embryo that originates from the fertilized egg cell, the triploid endosperm that is produced from the fertilized ...central cell, and the maternal sporophytic integuments that develop into the seed coat
. At the onset of embryo development in Arabidopsis thaliana, the zygote divides asymmetrically, producing a small apical embryonic cell and a larger basal cell that connects the embryo to the maternal tissue
. The coordinated and synchronous development of the embryo and the surrounding integuments, and the alignment of their growth axes, suggest communication between maternal tissues and the embryo. In contrast to animals, however, where a network of maternal factors that direct embryo patterning have been identified
, only a few maternal mutations have been described to affect embryo development in plants
. Early embryo patterning in Arabidopsis requires accumulation of the phytohormone auxin in the apical cell by directed transport from the suspensor
. However, the origin of this auxin has remained obscure. Here we investigate the source of auxin for early embryogenesis and provide evidence that the mother plant coordinates seed development by supplying auxin to the early embryo from the integuments of the ovule. We show that auxin response increases in ovules after fertilization, due to upregulated auxin biosynthesis in the integuments, and this maternally produced auxin is required for correct embryo development.
Somatic embryogenesis (SE), which is a process that involves the in vitro-induced embryogenic reprogramming of plant somatic cells, requires dynamic changes in the cell transcriptome. These changes ...are fine-tuned by many genetic and epigenetic factors, including posttranslational histone modifications such as histone acetylation. Antagonistically acting enzymes, histone acetyltransferases (HATs) and deacetylases (HDACs), which control histone acetylation in many developmental processes, are believed to control SE. However, the function of specific HAT/HDACs and the genes that are subjected to histone acetylation-mediated regulation during SE have yet to be revealed. Here, we present the global and gene-specific changes in histone acetylation in Arabidopsis explants that are undergoing SE. In the TSA (trichostatin A)-induced SE, we demonstrate that H3 and H4 acetylation might control the expression of the critical transcription factor (
) genes of a vital role in SE, including
,
(
),
(
) and
(
). Within the HATs and HDACs, which mainly positively regulate SE, we identified HDA19 as negatively affecting SE by regulating
,
and
. Finally, we provide some evidence on the role of HDA19 in the histone acetylation-mediated regulation of
during SE. Our results reveal an essential function of histone acetylation in the epigenetic mechanisms that control the
genes that play critical roles in the embryogenic reprogramming of plant somatic cells. The results implicate the complexity of Hac-related gene regulation in embryogenic induction and point to differences in the regulatory mechanisms that are involved in auxin- and TSA-induced SE.
Auxin contributes to almost every aspect of plant development and metabolism as well as the transport and signalling of auxin-shaped plant growth and morphogenesis in response to endo- and exogenous ...signals including stress conditions. Consistently with the common belief that auxin is a central trigger of developmental changes in plants, the auxin treatment of explants was reported to be an indispensable inducer of somatic embryogenesis (SE) in a large number of plant species. Treating in vitro-cultured tissue with auxins (primarily 2,4-dichlorophenoxyacetic acid, which is a synthetic auxin-like plant growth regulator) results in the extensive reprogramming of the somatic cell transcriptome, which involves the modulation of numerous SE-associated transcription factor genes (
s). A number of SE-modulated TFs that control auxin metabolism and signalling have been identified, and conversely, the regulators of the auxin-signalling pathway seem to control the SE-involved TFs. In turn, the different expression of the genes encoding the core components of the auxin-signalling pathway, the AUXIN/INDOLE-3-ACETIC ACIDs (Aux/IAAs) and AUXIN RESPONSE FACTORs (ARFs), was demonstrated to accompany SE induction. Thus, the extensive crosstalk between the hormones, in particular, auxin and the TFs, was revealed to play a central role in the SE-regulatory network. Accordingly, LEAFY COTYLEDON (LEC1 and LEC2), BABY BOOM (BBM), AGAMOUS-LIKE15 (AGL15) and WUSCHEL (WUS) were found to constitute the central part of the complex regulatory network that directs the somatic plant cell towards embryogenic development in response to auxin. The revealing picture shows a high degree of complexity of the regulatory relationships between the TFs of the SE-regulatory network, which involve direct and indirect interactions and regulatory feedback loops. This review examines the recent advances in studies on the auxin-controlled genetic network, which is involved in the mechanism of SE induction and focuses on the complex regulatory relationships between the down- and up-stream targets of the SE-regulatory TFs. In particular, the outcomes from investigations on Arabidopsis, which became a model plant in research on genetic control of SE, are presented.
The dataset contains the records of vascular plant species occurrences and distribution in old cemeteries (OC) of the Lower Dnipro region (Southern Ukraine). The analysed cemeteries were located in ...different types of landscapes (agricultural, rural and urban) and represent various ways of using their area (currently used, closed, abandoned). The floristic list includes 440 species of vascular plants (437
, 3
. The dataset demonstrates a sozological (Red-lists species) value of old cemeteries in Southern Ukraine. The cemeteries constitute refuges of native, rare and steppe flora and play a role of steppe habitat islands in a landscape almost completely transformed to arable land or urbanised.
This is the first dataset which contains information about flora of old cemeteries in Lower Dnipro region (Southern Ukraine). The dataset comprises 2118 occurrences of vascular plants (440 species) recorded in the years 2008-2021 in 13 old cemeteries of the Lower Dnipro region. The dataset includes information about 85 occurrences of rare species (23 species
, 3
) and 652 occurrences of 117 steppe species.