Shaded Environments Research Articles

Dactyliform appendage contributes to delayed selfing in the diminutive orchid Stigmatodactylus sikokianus

Societal Impact StatementUnderstanding plant reproductive strategies is vital for conserving endangered species. This study reveals a novel self‐pollination mechanism in the diminutive orchid Stigmatodactylus sikokianus, facilitated by the movement of a finger‐like appendage beneath the stigma. While this less‐than‐1‐mm‐long structure inspired the name Stigmatodactylus, its ecological function remained unknown for over 130 years—until now. The findings suggest that this appendage facilitates delayed self‐pollination, likely ensuring reproductive success under conditions of pollinator scarcity. Furthermore, the research highlights the value of integrating taxonomy with functional morphology to advance our understanding of pollination ecology.Summary Orchidaceae is one of the largest and most morphologically diverse plant families, largely due to intricate relationships with pollinators. However, autonomous self‐pollination is relatively common, likely as an adaptation to limited pollinator availability. This study examines the reproductive mechanism of the diminutive orchid Stigmatodactylus sikokianus, focusing on the finger‐like appendage beneath the stigma. The reproductive biology of S. sikokianus was investigated through pollination observations, artificial pollination experiments, and detailed studies of column morphology. Pollination experiments demonstrated that S. sikokianus primarily relies on autonomous self‐pollination, as indicated by high fruit set in bagged flowers and no fruit set in emasculated ones. Microscopic analysis revealed that the dactyliform appendage arcs toward the stigma, maintaining its straight structure, and eventually adheres to the stigma surface, allowing contact between the appendage apex and pollinia. UV microscopy showed thick bundles of pollen tubes penetrating the appendage adhered to the stigma and reaching the base of the column. The absence of pollen tube formation before contact between the appendage apex and pollinia suggests that autogamy occurs exclusively through the appendage at a later flowering stage. This study uncovers a novel self‐pollination mechanism in orchids, where the movement of the stigma appendage likely facilitates delayed selfing. This mechanism ensures seed production and likely reproductive success in S. sikokianus, enabling it to thrive in shaded environments with limited pollinator availability. The discovery highlights the value of integrating taxonomic and ecological approaches, as morphological traits provide valuable insights into species life histories.

Genome-Wide Identification of GmPIF Family and Regulatory Pathway Analysis of GmPIF3g in Different Temperature Environments.

Phytochrome-interacting factors (PIFs) play a crucial role in regulating plant growth and development. However, studies on soybean PIFs are limited. Here, we identified 22 GmPIF genes from the soybean genome and classified the GmPIF proteins into 13 subfamilies based on amino acid sequence homology, secondary and tertiary structures, protein structure, and conserved motifs. Genome-wide collinearity analysis revealed that fragment duplication events play a dominant role in expanding the GmPIF gene family. Cis-acting element analysis revealed that the GmPIF gene family is involved in light response, hormone response, biotic-abiotic stress response elements, and plant growth and development. Gene expression analysis in different temperature environments showed that the GmPIF family was found to be induced by phytohormone treatments, with a significant increase in the expression level of GmPIF3g. GmPIF3g plays a key role in the regulation of the entire network, and in addition, 30 proteins interacting with the GmPIF3g promoter were identified through the use of a novel biofilm interference technique. This technique showed that the transcription factor Dof (DNA binding with one finger) binds to the GmPIF3g promoter, and Y1H assays indicated that Dof regulates its expression by binding to the PIF promoter. These results provide a theoretical basis for further studies on the regulatory network of GmPIF genes to improve the structure of soybean plants under shade environments, as well as a new method for analyzing regulatory elements that interact with gene promoters.

Riverine songbirds capture high levels of atmospheric mercury pollution from brown food webs in forests by mercury isotopic evidence.

Elevated methylmercury (MeHg) exposure poses significant risks to bird health, behavior, and reproduction. Still, the risk of MeHg exposure to forest birds, accounting for over 80 % of the world's bird species, is poorly understood. This study combines Hg isotopes and video analysis, aiming to assess MeHg exposure risks to a forest riverine songbird, the spotted forktail (Enicurus maculatus) from a remote subtropical montane forest. Noticeably, 83 % of feather MeHg concentrations of adult forktails exceeded 5000 ng g-1, a threshold level potentially impacting bird reproduction, and 50 % of feather MeHg concentrations in forktail nestlings exceeded the threshold level of 1000 ng g-1, that potentially impacts the nestling growth. Forktail nestlings ingested ∼ 99 % of their MeHg from prey within brown food webs (i.e., from forest floor, aquatic, and emergent aquatic prey). The Hg isotopes reveal that MeHg along the bird food chain is mostly derived from in situ methylation of litterfall deposited atmospheric Hg0, with limited photo-demethylation (i.e., 4-12 %) in shaded forest environments. The risk of MeHg exposure of forest songbirds correlated positively with the proportion of prey consumed from brown food webs. We recommend incorporating resident riverine songbirds in monitoring programs to better evaluate the effectiveness of the Minamata Convention, especially in remote forest ecosystems where in situ MeHg production may be underestimated.

Alternative substrates and shading in the initial development of Croton lechleri seedlings

Croton lechleri (dragon's blood) is a forest species used by traditional Amazonian communities for medicinal purposes, with potential for use in degraded area recovery and as a source of additional income for some communities. However, silvicultural studies on this species are scarce. This study aimed to evaluate the influence of substrates and shading levels on seedling emergence and initial development of C. lechleri seedlings. A completely randomized statistical design with four replications was used, arranged in a 7 x 5 factorial setup (substrates and shading levels). The variables analyzed were seedling emergence, emergence speed index, relative emergence frequency, root and shoot length, root and shoot dry mass, stem diameter, and leaf count. Alternative substrates composed of carbonized rice husk and commercial substrate at all shading levels studied favored the emergence and emergence speed index of C. lechleri seedlings. In the 80% shade environment, the substrate consisting of forest topsoil + aged chicken manure (3:1) proved suitable for post-emergence seedling development of C. lechleri.

Integrated Transcriptomic and Metabolomic Analysis Reveals Possible Molecular Mechanisms of Leaf Growth and Development in Disanthus cercidifolius var. longipes.

Background:Disanthus cercidifolius var. longipes is an ancient relic plant unique to China. However, the typical shade-loving plant is largely exposed to the sun, which poses a major challenge to its conservation. Methods: This study explored dynamic changes in primary and secondary metabolites in D. cercidifolius leaves at different stages of development, combining metabolomics and transcriptome analysis to discuss the differentially accumulated metabolites (DAMs) and differentially expressed genes (DEGs). Results: The DAMs and DEGs were enriched in pathways related to photosynthesis, carbon (C) metabolism, anthocyanin synthesis, plant hormone signal transduction, and flavonoid synthesis. At the initial stage of leaf development, many primary metabolites were synthesized in the leaves. Before leaf maturity, many primary metabolites were converted into secondary metabolites. Combined transcriptome and metabolome analysis showed that the metabolites and genes related to anthocyanin synthesis and flavonoid metabolism were upregulated. In contrast, the genes related to C metabolism and C fixation were downregulated. After leaf maturity, photosynthetic capacity increased, total flavonoid content peaked (implying the strongest photoprotection capacity), and the transformation of anthocyanins and flavonoids was weakened. Conclusions: Light intensity indirectly affects the accumulation of the primary and secondary metabolism of D. cercidifolius. With the enhancement of photoprotection, the photosynthetic energy capacity decreases. It is, therefore, inferable that D. cercidifolius has shading properties and achieves a stable nutrient supply during growth and development through these strategies. Thus, D. cercidifolius protection requires a shaded environment.

Advancements in Cardamom (<i>Elettaria cardamomum </i>Maton) Breeding: Genetic Diversity, Biotech Strategies, and Future Directions

Cardamom (<i>Elettaria cardamomum </i>Maton), often referred to as the 'Queen of Spices,' belongs to the Zingiberaceae family and is noted for its unique flavor and aroma. As an herbaceous perennial, cardamom thrives in the shaded environments of the Western Ghats in South India, where it exhibits significant genetic diversity necessary for breeding improvement. This paper explores the genetic resources of cardamom, including advances in breeding methodologies and biotechnological approaches aimed at enhancing productivity and quality. Key breeding strategies include clonal selection, hybridization, and mutation breeding, which have led to the development of several high-yielding and stress-resistant varieties. The integration of molecular techniques, such as marker-assisted selection and genetic transformation, offers prospects for tapping into cardamom's genetic diversity for future improvements. However, the genetic erosion of natural cardamom populations necessitates urgent conservation efforts to safeguard this valuable germplasm. This review emphasizes the importance of innovative breeding techniques and strategic direction for the sustainable enhancement of cardamom as a critical global spice crop.

Cultivation conditions and their impact on yerba mate ecophysiology and antioxidant potential

This work aimed to evaluate the effect of light and shading on yerba mate's ecophysiology and antioxidant potential. To this end, an experiment was carried out in an entirely randomized design, with three treatments, being cultivated in full sun (Ts), in agroforestry (Ta), and in native forest (Tm). Photosynthesis activity and chlorophyll, phenolic compounds, flavonoids, and tannins , and antioxidant activity were analyzed to determine the best cultivation environment that influences the ecophysiology and antioxidant potential of Yerba mate. Ts environment yielded higher concentrations of phenolic compounds, flavonoids, and tannins. Shaded environments showed greater antioxidant activity. Chlorophyll and photosynthesis indices differed in shaded environments, with Tm having higher concentration and photosynthetic activity and Ta having lower concentration and photosynthetic activity. The results show that PAR positively influenced the biochemical parameters. Shading was more efficient for antioxidant activity, and agroforestry contributed to greater antioxidant activity. The highest concentrations of chlorophyll and highest photosynthetic activity were found in yerba mate in native forests.

Sulfur affects multi-scale starch structures and its contribution to the cookie-baking quality of wheat subjected to shade stress

The components and structure of starch macromolecules critically determine its food-use properties. However, elemental sulfur supplementation affects the relationship between starch structure and the cookie-making quality of wheat under shaded environments remains unclear. Here, we investigated the effect of sulfur on the starch multi-scale structures and its contribution to the cookie-baking quality of wheat after pre- or post-anthesis shading. Compared with the unshaded control, shade stress decreased the amylose and total starch contents, formed smaller B-type starch granules, narrowed the molecular weight distribution, and decreased the amylopectin long-chain proportion, crystallinity, viscosity, and spread ratio of cookies. Weak-gluten cultivars are more sensitive to shade stress than strong-gluten cultivars. Under shaded environments, sulfur increased the amylopectin content, proportion of amylopectin short chains, and total starch content, increasing the mean diameter of starch granules and viscosity, ultimately decreasing the cookie hardness. The random forest model revealed that the surface area of the starch granules (18.7 %) and amylopectin B3 chain (6.7 %) contributed the most to the variation in the cookie spread ratio. Cookie hardness was determined mainly by the total starch (7.8 %), amylopectin (6.3 %), and trough viscosity (5.0 %). Our results help to design strategies for achieving superior-quality wheat in the context of global dimming.

Quality of coffee dried on suspended terrace with different layers

Seeking to improve the final quality of coffee, producers began to improve and innovate in all stages of this segment to meet the demanding market. Hence, this study evaluated the physicochemical and sensory quality of dried unripe, cherry and over-ripe coffee cherries on suspended terraces in different layers. The experiment involved drying coffee in all layers of the suspended terraces, consisting of seven treatments and four replications as follows: T1– concrete terrace; T2 – conventional “single-layer” suspended terrace; T3 – first layer “suspended terrace with two layers”; T4 – second layer “suspended terrace with two layers”; T5 – first layer “suspended terrace with three layers”; T6 – second layer “suspended terrace with three layers”; and T7 – third layer “suspended terrace with three layers.” All treatments used identical samples composed of a mixture of coffee cherries in the three different physiological maturation stages (unripe, cherry and over-ripe). Percentage of each stage was evaluated in a one-liter container with coffee. Physicochemical and sensory quality of unripe, cherry and overripe coffees is influenced by the terrace structure. Treatments T5 and T6, represented by the first layer “suspended terrace with three layers” and second layer “suspended terrace with three layers,” presented the best results for the beneficial fermentation caused by the terrace structure, providing a more shaded environment and resulting in desirable sensory attributes.

The effect of phosphorus, irradiance and competitor identity on the relative performance of invasive Chromolaena odorata

BackgroundResource competition is an important factor affecting the invasion success of alien plants, and environmental factors influence the competition outcomes between invasive and native plants. In this study, we explore the competitive pattern between invasive Chromolaena odorata and two native plant species under different phosphorus and irradiance levels.ResultsThe final biomass of each plant was regulated by both morphological and physiological traits. Invasive C. odorata did not always perform better than both native plants, and the competitive pattern between C. odorata and native plants was dependent on native competitor identity and environmental conditions. With competition, invasive C. odorata showed higher biomass (over 60%) than native Xanthium sibiricum under all treatments, but only showed higher biomass (about 20%) than native Eupatorium lindleyanum in normal irradiance treatments. The effect of phosphorus on competition depended on the irradiance level. Under normal irradiance, phosphorus addition increased (almost 10 times) the competitive index of invasive C. odorata; however, under shade irradiance, phosphorus addition decreased (40%) the competitive index of C. odorata.ConclusionThese results suggest that phosphorus, irradiance and native plant competitor together influence the relative performance of invasive C. odorata. In shade environment, selecting E. lindleyanum as competitor and increasing phosphorus level is an effective method for controlling the invasion of C. odorata.

Initial stomatal conductance increases photosynthetic induction of trees leaves more from sunlit than from shaded environments: a meta-analysis.

It has long been held that tree species/leaves from shaded environments show faster rate of photosynthetic induction than species/leaves from sunlit environments, but the evidence so far is conflicting and the underlying mechanisms are still under debate. To address the debate, we compiled a dataset for 87 tree species and compared the initial increasing slope during the first 2-min induction (SA) and stomatal and biochemical characteristics between sun and shade species from the same study, and those between sun and shade leaves within the same species. In 77% of between-species comparisons, the species with high steady-state photosynthetic rate in the high light (Af) exhibited a larger SA than the species with low Af. In 67% within-species comparisons, the sun leaves exhibited a larger SA than the shade leaves. However, in only a few instances did the sun species/leaves more rapidly achieve 50% of full induction, with an even smaller SA, than the shade species/leaves. At both the species and leaf level, SA increased with increasing initial stomatal conductance before induction (gsi). Despite exhibiting reduced intrinsic water-use efficiency in low light, a large SA proportionally enhances photosynthetic carbon gain during the first 2-min induction in the sun species and leaves. Thus, in terms of the increase in absolute rate of photosynthesis, tree species/leaves from sunlit environments display faster photosynthetic induction responses than those from shaded environments. Our results call for re-consideration of contrasting photosynthetic strategies in photosynthetic adaption/acclimation to dynamic light environments across species.

Different habitat condition proportions on farms affect the structure and diversity of dung beetle (Coleoptera, Scarabaeidae, Scarabaeinae) communities

The continuous changes and increase of land use into ranching and agriculture have caused shifts in the composition and structure of dung beetle communities due to the modification of vegetation structure. The impact of these changes can be approached using dung beetles as ecological indicators. Agricultural, ranching, and forestry are often integrated into the same farm in different proportions (landscape level), and their degree of integration with habitats changes over time and space. We assessed if different habitat proportions of crop, pasture, and forest land on three farms affect the structure and diversity of dung beetle (Coleoptera: Scarabaeinae) communities. Farms included from three to four habitat conditions each (plantations of timber species, banana-coffee plantation, living fences, pastures, secondary and riparian forest). Pitfall traps with dung and carrion as bait were placed across each habitat condition of each farm during three different sampling periods. Across all samplings, 1,198 dung beetle individuals belonging to 21 species were captured. Species diversity and composition vary according to the type of farm and the main factor that modifies this tendency is the proportion of pasture land composing the farm. Farms with a lower proportion of grazing land (11.96% for forestry farms and 32.19% for agricultural farms) and denser vegetation canopy, which cast more shadows, had greater beetle diversity. Conversely, farms having a greater proportion of grazing land had low diversity and a dominant tendency in the species abundance curve. Umbraphile species dominated the forest farm, indicating a strong affinity for shaded environments such as the one provided there, while species displaying no habitat preference exhibited higher abundance in the ranching farm. Additionally, a notable prevalence of small-sized species was observed in the ranching farm, contrasting with a relatively even distribution of sizes in the remaining landscapes. These tendencies suggest that shade positively influences biodiversity conservation. Riparian vegetation, living fences, and banana-coffee plantations are important connectivity elements in agricultural landscapes for shade-adapted dung beetle species.

A star selection strategy based on MDS and fast convex hull algorithm in COSPAS-SARSAT system

Abstract The space segment of the current COSPAS-SARSAT system is mainly supported by the Galileo system, and the positioning technology used is generally three-star time-difference positioning. Most of the antennas equipped in the Mission Control Centre (MCC) are not omnidirectional. In the case of Source of Search and Rescue (SAR) in a heavily shaded environment, it is necessary to use the different orbit time-sharing single satellite geolocations using TDOA, and the time-sharing will generate more satellite position data. In order to achieve the higher positioning accuracy of the solution, it is necessary to select the appropriate time and optimal geometrical distribution from the satellite positions. In this paper, we propose a satellite selection strategy based on Multiple Dimensional scaling (MDS) data degradation and a fast convex hull algorithm. The three-dimensional satellite data are downscaled to two-dimensional while maintaining their relative positional relationships, and the two-dimensional convex hull algorithm is used for Geometric Dilution of Precision (GDOP) minimum search star selection. By simulating the satellite position data in different orbits and time-sharing to complete the localization solution, the results show that compared with the traditional satellite selection method, the satellite selection strategy proposed in this paper is able to achieve better localization accuracy in a shorter period of time.

A Holistic Approach to the Selection of Soybean (Glycine max) Cultivars for Shade Environments Based on Morphological, Yield and Genetic Traits

ABSTRACTThe reduction of photosynthetically active radiation impacts the growth and productivity of soybean in agroforestry and intercropping systems. Thus, this report explored the responses of 16 soybean cultivars submitted to shade levels in field conditions. Multi‐faceted and relative importance analyses revealed that the steam diameter and plant height are fundamental morphological markers for selecting shade‐resilient cultivars, both were high and positively correlated to yield components. Moreover, the responses to shade varied among soybean cultivars, with certain genotypes demonstrating distinct tolerance levels, which allowed also the estimative of genetic variance that revealed strong participation of genetic components in responses to shade. Multivariate and clustering analysis using steam diameter and plant height in combination with two yield components resulted in the identification of four soybean cultivars more tolerant to shade environments and two sensible. Therefore, this report provides insights into soybean cultivation under varying light conditions, provides a robust foundation for the integration of morphological and yield markers in breeding programmes focused on shade tolerance and guides future endeavours in crop improvement for optimal and sustainable yield and resilience in the climate change context.

Differences in anterior segment structure and pupillary diastolic characteristics of eyes with primary angle closure suspects under light and shade environments

Objective: To investigate the anatomical structure changes of the anterior segment and dynamic pupil changes in eyes with suspected primary angle-closure (PACS) under light and dark conditions, and their correlation with the occurrence of acute primary angle-closure (APAC). Methods: This cross-sectional study collected data from 37 PACS patients (66 eyes) who visited the ophthalmology clinic of the Second Affiliated Hospital of Harbin Medical University between September 2019 and March 2021. The patients included 12 males and 25 females, with an average age of (61.27±7.35) years. Of the 66 eyes, 58 had no history of APAC in the contralateral eye, while 8 had a history of APAC in the contralateral eye. Patients without a history of APAC in both eyes underwent the dark room prone provocative test (DRPPT), and the eyes were divided into three groups: DRPPT- (44 eyes), DRPPT+ (14 eyes), and APAC (8 eyes). The DRPPT+ and APAC groups were combined into the APAC+ group. All patients underwent ultrasound biomicroscopy (UBM) to measure the changes in various parameters under light and dark conditions, including anterior chamber angle 500 (ACA500Δ) and 1000 (ACA1000Δ), angle opening distance 500 (AOD500Δ) and 1000 (AOD1000Δ), and iris thickness 500 (IT500Δ) and 1000 (IT1000Δ). Anterior segment analysis was performed to capture the pupil changes under light and dark conditions, recording pupil diameter, maximum dilation speed (Vmax), maximum constriction speed (Vmin), and average speed (Vm). Results: There was no significant difference in the parameters between DRPPT+ group and APAC group (P>0.05). In the difference analysis, it was found that the Vm value of DRPPT- group [(0.17±0.07) mm/s] was higher than that of APAC+ group [(0.13±0.06)mm/s], and the difference was statistically significant (P<0.05). There was no significant difference in other parameters (P>0.05). Vmax was positively correlated with temporal ACA1000Δ and temporal AOD1000Δ in all patients with PACS (r=0.302, 0.260; P<0.05), Vmin was negatively correlated with temporal ACA1000Δ and temporal AOD1000Δ (r=-0.338, -0.330; P<0.05). Conclusions: The dynamic changes in the anterior segment and pupil under different lighting conditions provide insights into the risk factors and potential predictive indicators for the occurrence of APAC in PACS patients.

Innovative GMPPT searching algorithm and precise backstepping control for grid-connected PV system in challenging shading environments

Innovative GMPPT searching algorithm and precise backstepping control for grid-connected PV system in challenging shading environments

How does the life cycle of Clinodiplosis profusa (Cecidomyiidae) adjust to phenological variations of the host plant Eugenia uniflora (Myrtaceae) in sun and shade?

Galls are plant neoformations induced by specialized parasites. Since gall inducers rely on reactive plant sites for gall development, variations in abiotic factors that affect plant phenology are expected to impact the life cycle of gall inducers. To test the hypothesis that different light conditions affect both host plant and gall inducer life cycles, we studied the system Eugenia uniflora (Myrtaceae) - Clinodiplosis profusa (Cecidomyiidae), comparing plants occurring in sunny and shaded environments. We mapped phenological differences among individuals of E. uniflora occurring in the two environments and related them to the influence of luminosity on the life cycle of the gall inducer. Shade plants showed lower intensity of leaf sprouting throughout the year compared to sun-exposed plants, especially during the rainy season. Young and mature galls are synchronized with the peak of leaf sprouting at the beginning of the rainy season, lasting longer in sun-exposed plants - approximately two months longer compared to shade plants. The greater light intensity positively impacts the formation and growth of leaves and galls, with an extended period available for their induction and growth. Thus, light is an important factor for the development of gallers, considering that variations in luminosity influenced not only the phenology of the host plant, but also determined the life cycle of gall inducers. Furthermore, changes in plant-environment interactions are expected to affect the life cycle and richness of other host plant-gall inducer systems.

Ontogenetic differences in sun and shade galls of Clinodiplosis profusa on Eugenia uniflora leaves and the cytological antioxidant mechanisms in gall cells.

The gall-host Eugenia uniflora (Myrtaceae) is adaptable to different light conditions, enabling leaf production and survival in both sun and shade. Leaves of E. uniflora in shaded environments have more mesophyll layers, and galls of Clinodiplosis profusa (Cecidomyiidae) are larger and wider. Based on these previous observations, this study investigated the morphogenesis of galls induced by C. profusa on leaves of E. uniflora in different light conditions, revealing if the galls have a potential for acclimation, as observed with leaves. For this purpose, we compared the anatomical, histometric, and histochemical development of leaves and galls at different stages of development in sun and shade environments. Additionally, we analyzed the cytological features of the tissues composing the mature gall walls. Cells of shade galls expanded more toward the end of the developmental phase, which may explain the larger volume found for shade galls in a previous study. However, during the mature phase, these galls showed no significant differences in tissue thickness and final cell elongation in the contrasting light conditions. In the ultrastructural analyses, mature galls showed a gradient distinguishing the outer and inner parenchyma cells. The inner parenchyma had nutritive cells, with dense cytoplasm and abundant organelles. A higher accumulation of starch grains in nutritive cells, with evidence of hydrolysis of starch grains detected in the innermost layers leads to the accumulation of reducing sugars, which, with the presence of plastoglobules and protein bodies, are important mechanisms of oxidative stress dissipation in the cells in contact with the gall inducer.

Protected cultivation and water salinity in the formation of Erythrina velutina Willd seedlings

Protected cultivation can reduce the harmful effects of salt stress on the development of plant species. The objective of this study was to evaluate the growth and morphophysiological responses of E. velutina seedlings grown under saline stress in two growing environments. The experiment was conducted in an experimental area belonging to the Federal University of Ceará, in Fortaleza, Ceará. The experimental design was entirely randomized in subdivided plots, where the plots were represented by the growing environments (closed-screen greenhouse - CSG and open-sided greenhouse chapel - OSG), the subplots by the irrigation waters (0.8 and 5.0 dS m-1) and in the subplots the collection seasons (20, 40, 60 and 80 DAS), with four replications. Saline stress inhibited the growth of E. velutina seedlings in terms of height, leaf area, number of leaves and dry mass of the aerial part, but provided greater root dry mass throughout the growing season. It also affected gas exchange after 50 days of cultivation. The seedlings grown in the open shade environment on the sides and with lower salinity water had a higher Dickson quality index.

Bee pollination in the shadows: The role of pseudopollen and agamospermy in the mycoheterotrophic orchid Gastrodia elata

Societal Impact StatementThe family Orchidaceae is renowned for its reliance on specialized pollinators and mycorrhizal symbioses. This study examined how the nonphotosynthetic orchid Gastrodia elata thrives on the dim forest floor, focusing on its pollination biology, particularly the chemical properties of pseudopollen and reproductive assurance mechanisms. It was found that G. elata mainly relies on Lasioglossum bees, attracted by its starch‐rich pseudopollen. Additionally, G. elata in the studied population can produce fruit through agamospermy (seeds from unfertilized ovules). Overall, the findings reveal that G. elata relies on fungi for nutrition and employs reproductive strategies such as mimicking pollen to attract bees and resorting to asexual reproduction when pollinators are scarce.Summary Mycoheterotrophy is an adaptation that allows survival in environments with low irradiance and minimal competition from autotrophic plants. However, such environments can negatively impact bee pollination, as most bees prefer open habitats. Despite this, the mycoheterotrophic orchid Gastrodia elata is known to rely on Lasioglossum bees for pollination. We investigated the reproductive biology of G. elata to understand how it overcomes pollinator limitation. Our research focused on the chemical composition and anatomical basis of G. elata pseudopollen (a substance that mimics pollen and may facilitate pollination), based on micromorphological and nutritional analyses, as well as observations of pollinator behavior. Additionally, we explored the potential presence of autogamy or agamospermy as mechanisms for reproductive assurance. The pseudopollen in G. elata originates from the disintegration of the adaxial parenchymatous tissue of the callus and is rich in starch. This contrasts with pseudopollen in most orchids, which typically originates from lip hairs and whose potential attractant is protein. Lasioglossum bees pollinating G. elata visit multiple flowers and actively collect pseudopollen. Furthermore, agamospermy serves as a reproductive safeguard in shaded habitats where insect‐mediated pollination is infrequent, at least in the investigated population. Our findings suggest that Gastrodia elata pseudopollen plays an important role in enhancing insect‐mediated pollination, particularly in relatively open habitats, while agamospermy ensures fruit set in shaded environments of the investigated population. Both pseudopollen production and agamospermy likely help overcome the pollination constraints posed by the mycoheterotrophic lifestyle.