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Lamina traits and structure evaluated in transverse sections of Eugenia hiemalis first leaves. Leaves developed in 100 % (a), 50 % (b), 30 % (c), and 0.3 % (d) of full sunlight (mean ± SE). Different letters indicate significant differences of mean values (P < 0.05) among the treatments. Bars 50 μm
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Leaf light responses, especially structural aspects, are well studied in adult tropical tree species. However, there is still a gap in the knowledge of these plastic responses on first developed leaves, which may be important to seedling survival in variable environments. An experimental approach was used to test the hypothesis that phenotypic plas...
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Background and aims:
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Citations
... Particularly, as light intensity decreases, leaf thickness, thickness of palisade and spongy tissues decrease, and the number of palisade tissue layers decreases (Putz and Mooney, 1991), which are adaptations of plants to low light environments. For example, Nascimento et al. (2015) revealed that leaf and mesophyll thickness was greater under high light intensity in Eugenia hiemalis. LSs is significantly thinner than LFs, and it also has fewer palisade and spongy tissues, suggesting it adapts to low light environments. ...
Introduction
Plants that display heteroblasty possess conspicuous variations in leaf morphology between their juvenile and adult phases, with certain species retaining juvenile-like leaves even in adulthood. Nevertheless, the ecological advantages of maintaining two or more distinct leaf types in heteroblastic plants at the adult stage remain unclear.
Method
The aim of this study is to examine the adaptive significance of heteroblastic leaves sampled from branches with divergent functions (sterile and fertile branches) of mature Ficus pumila individuals by comparing their morphological, anatomical, and physiological characteristics.
Result
Leaves on sterile branches (LSs) exhibited a significantly larger specific leaf area, thinner palisade and spongy tissues, lower chlorophyll contents, and lower light saturation points than leaves on fertile branches (LFs). These results demonstrate that LSs are better adapted to low light environments, while LFs are well equipped to take advantages of high light conditions. However, both LFs and LSs have a low light compensation point with no significant difference between them, indicating that they start to accumulate photosynthetic products under similar light conditions. Interestingly, significant higher net photosynthetic rate was detected in LFs, showing they have higher photosynthetic capacity. Furthermore, LFs produced significant more nutrients compared to LSs, which may associate to their ability of accumulating more photosynthetic products under full light conditions and higher photosynthetic capacity.
Discussion
Overall, we observed a pattern of divergence in morphological features of leaves on two functional branches. Anatomical and physiological features indicate that LFs have an advantage in varied light conditions, providing amounts of photosynthetic products to support the sexual reproduction, while LSs adapt to low light environments. Our findings provide evidence that heteroblasty facilitates F. pumila to utilize varying light environments, likely associated with its growth form as a climbing plant. This strategy allows the plant to allocate resources more effectively and optimize its overall fitness.
... The effects of light on the leaf epidermis are also well known, including commonly reduced stomatal density in shade leaves compared to sun leaves due to the greater size of epidermal cells and stomata (Dickinson, 2000;Nascimento et al., 2015;Pompelli et al., 2010). In some cases, only cell hypertrophy leads to reduced stomatal density (Pompelli et al., 2010). ...
... Although the mesophyll features exhibited distinct responses compared to other plants, the stomatal density and epidermal cell sizes had responses common to shade and sun leaves of other plant species, including Eugenia spp. (Donato and Morretes, 2009;Lemos et al., 2018Lemos et al., , 2019Nascimento et al., 2015). The distinct responses of the current studied plants to sunlight are also indicative of the better adaptation of such plants to xeric than mesic conditions, and the plasticity of epidermal features compensates for the differences in sunlight exposure by finely controlling water transpiration rates and CO 2 assimilation (Lemos et al., 2018(Lemos et al., , 2019Nascimento et al., 2015). ...
... (Donato and Morretes, 2009;Lemos et al., 2018Lemos et al., , 2019Nascimento et al., 2015). The distinct responses of the current studied plants to sunlight are also indicative of the better adaptation of such plants to xeric than mesic conditions, and the plasticity of epidermal features compensates for the differences in sunlight exposure by finely controlling water transpiration rates and CO 2 assimilation (Lemos et al., 2018(Lemos et al., , 2019Nascimento et al., 2015). Distinct responses were also observed in sun and shade leaves of Olea europaea, which is influenced by polymorphisms and responses to diffuse light (De Casas et al., 2011). ...
Gall-inducing insects stimulate their host plants, leading to the formation of distinct structures, such as the fusiform galls induced by Clinodiplosis profusa Maia, 2001 (Cecidomyiidae) on leaves of Eugenia uniflora L. (Myrtaceae). The influence of different light conditions on infestation rates, establishment, and morphogenesis of galls is little known. We tested if gall infestation is higher in E. uniflora sun plants compared to shade plants, and if the size, anatomical, and histometrical features of the mesophyll and epidermis are distinct between sun and shade leaves and galls. Gall infestation rates are similar in sun and shade plants, although the results indicate a possible difference in life cycle duration between the light conditions. Anatomical data revealed the leaves and galls respond similarly to sun and shade for number of cell layers. In the galls the hyperplasia provoked by C. profusa is influenced by distinct sunlight exposure. Cell hypertrophy and a reduced stomatal index in the galls is remarkable when compared to non-galled leaves. The cell sizes in the mesophyll and epidermis and stomatal densities are similar between the sun and shade galls, but these factors are distinct between the sun and shade leaves. Cell hypertrophy and epidermal differentiation patterns are more influenced by the gall inducer, which inhibits the influence of light on these patterns. In conclusion, gall morphogenesis is strongly influenced by galling stimuli for some developmental processes, but other processes are influenced by abiotic conditions, such as light exposure. This opens a field in gall ecological anatomy and development.
... in relation to spatial-scale environmental heterogeneity [7][8][9], much less attention has been paid to the relationship between time-scale environmental heterogeneity and plasticity. However, temporally heterogeneous environments may be more ubiquitous than environmental heterogeneity at spatial scale, plants can experience fluctuations in light, water and nutrient availability at scales of hours, days and months in their lifetime It is reported that Convolvulus chilensis experiencing the greatest temporal variation in nature has shown the greatest plasticity in four traits [10]. ...
Temporally heterogeneous environments is hypothesized to correlate with greater plasticity of plants, which has rarely been supported by direct evidence. To address this issue, we subjected three species from different ranges of habitats to a first round of alternating full light and heavy shading (temporally heterogeneous light experience), constant moderate shading and full light conditions (temporally homogeneous light experiences, control) and a second round of light-gradient treatments. We measured plant performance in a series of morphological, biomass, physiological and biochemical traits at the end of each round. Compared to constant full light experience, temporally heterogeneous light conditions induced immediate active biochemical responses (in the first round) with improved late growth in biomass (during the second round); constant moderate shading experience increased photosynthetic physiological and biomass performances of plants in early response, and decreased their late growth in biomass. The karst endemic species of Kmeria septentrionalis showed greater improvement in late growth of biomass and lower decrease in biochemical performance, due to early heterogeneous experience, compared to the non-karst species of Lithocarpus glaber and the karst adaptable species of Celtis sinensis. Results suggested plants will prefer to produce morphological and physiological responses that are less reversible and more costly in the face of more reliable environmental cues at early stage in spite of decreased future growth potential, but to produce immediate biochemical responses for higher late growth potential when early environmental cues are less reliable, to avoid the loss of high costs and low profits. Typical karst species should be more able to benefit from early temporally heterogeneous experience, due to long-term adaptation to karst habitats of high environmental heterogeneity and low resource availability.
... Moreover, the leaves of introduced species compared to the native seem to have a high plastic response to light levels (Yamashita et al., 2000). Nascimento et al. (2015) suggested that functional traits that play critical roles in survival and development are susceptible to change and adapt to different environments. Considering these studies, L. lucidum has invaded coastal areas by changing stomatal size, suggesting that L. lucidum may have the capacity for acclimation owing to phenotypic plasticity. ...
Alien, introduced, and non-native species expand their distribution through the development of various transportation networks. In Japan, Ligustrum lucidum Aiton (Oleaceae) was introduced in the mid-1800s for use as roadside trees, but has escaped from the planting area and spread everywhere. This species also invades coastal areas and has adapted to develop its characteristic features under various stress conditions. To clarify the adaptive features of L. lucidum in coastal areas, we conducted morphological and anatomical analyses. We found that the stomatal size of L. lucidum was smaller in the coastal populations than in inland populations, which is similar to the coastal adaptation pattern of L. japonicum Thunb., a closely related native species. Our results suggest that strong selective pressure on coastal area conditions could force their leaves to avoid excessive transpiration, and such an adaptation pattern of L. lucidum could expand its distribution to various coastal areas.
... High values have been reported in invasive woody species (Grotkopp and Rejmanek 2007). IAS when compared to non invasive plant species seem to have a high plastic response to light levels (Yamashita et al. 2000) which plays a key role in plant survival and development (Nascimento et al. 2015). Further, many recent studies revealed that reproductive dispersion traits such as seed dry mass, seed dispersal method, seed production and plant maximum height are highly correlated with biological invasion (Moodley et al. 2013;Moravcova et al. 2015). ...
Invasive alien species (IAS) poses a significant threat to plant biodiversity globally and even considered one of the largest threats to biodiversity, second to habitat loss. They behave as pioneer species in different landscapes, tolerant to disturbances, climatic conditions, high competitive potential and generalists in distribution. Their superior competitive ability results in the loss of native flora leading to extinction. The success of IAS generally attributed to differences in functional traits compared to less successful aliens as well as to native species. Several studies envisaged that the impacts of plant invasions are not universal and depend on the trait diversity of both, the introduced species and the resident community. Functional traits best describe the alien's success over natives , and they seem to be important attributes to conservation biology and ecosystem management. Moreover, their ecological impacts remain poorly understood due to lack of quantitative studies. In the present paper, we adopted the systematic literature review approach for collecting and analysing the scientific data. A total of 212 critical research papers and grey literature for last three decades were found meeting the aims, were collected from relevant sources. Present review emphasizes the differences in key functional traits between invasive alien plants and the native species which aid them to alter ecosystem functioning by modifying habitat according to their needs. We also focus on potential habitats for invasion based on a conceptual framework concerning response-effect traits. Review provides a quantitative assessment of invading species for their ecological performance , emerging problems and possible solution.
... The light environment varies considerably during forest development and succession at both temporal and spatial scales [1][2][3]. This phenomenon hampers or accelerates plant growth intermittently, especially beneath the forest canopy [4]. ...
Effects of light quality on phenotypic plasticity in Cunninghamia lanceolata (Lamb.) Hook.
seedlings during growth and development, and the underlying mechanisms, were investigated. The seedlings showed distinct morphological adjustments when exposed to an equal photosynthetic photon flux density (400 �mol�m2�s1) of different light qualities: monochromatic blue (BL), monochromatic red (RL), monochromatic far-red (FrL), mixed RL and FrL at 1:1 (RFr1:1L), mixed RL and FrL at 1:2 (RFr1:2L), and multi-wavelength white (WL, control). Compared with WL, FrL and BL significantly promoted height increment. However, BL was unfavorable for root growth. The seedling biomass was lower and the root-to-shoot ratio was smaller under BL. RL promoted leaf area enlargement, root growth, axillary bud number, and increased the root-to-shoot ratio, but inhibited stem elongation. Low R/Fr ratios or increased FrL proportion increased seedling stem elongation. The seedling growth under RFr1:1L treatment was poorer than that under other treatments; however, the number of axillary buds was the highest. The plasticity of leaf morphology traits was lower in different treatments, and that of axillary bud traits was crucial in the adaptation of C. lanceolata to light quality. Precise management of light quality and wavelength in controlled environments may maximize the economic efficiency of forest production and enhance its quality.
... Because excessive irradiance is detrimental to photosynthetic tissues, plants must produce smaller and thicker leaves with a higher leaf mass per unit area under intense light conditions. This morphology of C. lanceolata leaves allows heat dissipation and helps avoid damage from overheating and high transpiration rates [35,36]. ...
The improvement of the stand yield and economic benefits of Cunninghamialanceolata (Lamb.) Hook, one of the most important plantation trees in China, has always been a concern. An appropriate planting density plays an important role in increasing productivity. To determine the optimum planting density of C. lanceolata, we measured leaf morphology, gas exchange parameters, and photosynthetic pigments in replicate leaves on 4-year-old saplings grown in four canopy positions under a low (D1), medium (D2), and high (D3) planting density (approximately 900, 2505, and 6660 trees·ha−1, respectively). We then ranked trait variations using the coefficient of variation to explore the influence of planting density. Planting density significantly influenced the leaf morphology, gas exchange parameters, and the photosynthetic pigment contents of C. lanceolata. Medium planting density (D2) resulted in a larger leaf area and specific leaf areas, a higher net photosynthesis rate, and higher photosynthetic pigment contents. The responses of leaf gas exchange parameters had the most plastic traits that responded to planting density. An appropriate planting density and leaf position might improve C. lanceolata leaf morphology and physiological attributes, which in turn would facilitate growth.
... Species with broad geographic distribution in heterogeneous environments, Eugenia uniflora and E. hiemalis, displayed wider leaves, whereas E. subterminalis and E. pyriformis, which are more restricted, presented narrower leaves. In fact, a high phenotypic plasticity had already been detected in E. hiemalis, both at the individual and population levels (Nascimento et al., 2015). According to Gates (1980), larger and wider leaves (E. ...
... subterminalis and E. pyriformis). Thus, the observed differences in leaf shape may indicate tolerance to lighted environments, which has already been observed for Eugenia by Cardoso and Lomônaco (2003) and associated with phenotypic plasticity (Nascimento et al., 2015). ...
Eugenia (Myrtaceae) is one of the most diverse groups in Neotropical flora. This study evaluated leaf size and shape in four species of Eugenia as a model to test geometric morphometrics in species with distinct geographic distribution in Atlantic Forest biome. The species E. hiemalis Cambess, E. pyriformis Cambess, E. subterminalis DC and E. uniflora L. were selected and the leaves used were analysed from herbariums and scientific collections. Leaves were photographed from a standardized distance and anatomical landmarks and semi-landmarks were placed on the contour of each leaf. The leaf centroid size, leaf shape and the phylogenetic signal were analysed. The phylogenetic signal was obtained from tree available in literature for Eugenia genus. Leaf size differed amongst species. E. pyriformis and E. hiemalis displayed a larger leaf centroid size when compared to E. uniflora and E. subterminalis. For the leaf shape was observed that E. pyriformis and E. subterminalis present narrower leaves, different from E. hiemalis and E. uniflora with wider leaves. There is no phylogenetic signal in leaf morphology among the four species. The results indicate that species with broad geographical distribution or associated with areas of high environmental heterogeneity resulting in high leaf form amplitude. Phenotypic plasticity could be detected by geometric morphometrics, indicating this tool as a useful approach to quantify leaf shape variation in Eugenia, at least in these studied species.
... Seedling growth and development are critical to population dynamics, and they include complex processes of detecting, acceptance, and interpreting endogenous and environmental signals [1]. Plants respond to endogenous and environmental signals in many different ways [2]. ...
... Among the four treatments, one treatment was 100% natural light photon irradiance (bare land), and different shades were then adjusted by covering three different densities of specifically shaded cloth on three iron frames (Meshel Netting Co., Ltd, Changzhou, Jiangsu, CN; customized one needle, two needle, and four needle three specifications of shading cloth by the company, it spectrally provides rather uniform shade at those levels of shade presently marketed) with a height of 2.0 m. For the photon radiation gradient, the shading cloth was 20 cm away from the ground to ensure ventilation [1,33]. The seedlings were randomly divided into four groups and placed in the aforementioned regions with four light treatments. ...
... This is consistent with the previous research results. Under full light, the thickness of the plant leaf increases, epidermis structure is developed, and palisade tissue is developed [1,42], whereas leaf thinning, spongy tissue porosity, and ventilation are adaptive characteristics under low light conditions [43]. Under full light, plants retain their water contents by having thick mesophyll tissue, thick epidermis, and strong refraction, which effectively regulates the mesophyll tissue protecting it from damage from strong light [44,45]. ...
This study examined the effects of light intensity on the plasticity of the leaves of Juglans regia f. luodianense seedlings in karst habitat and how they respond to changes in light intensity. The light intensity of 1-year-old seedlings of J. regia f. luodianense in different niches in a karst area was set as 100% (bare land), 75% (forest margin), 50% (forest gap), and 25% (under forest) of natural light. The material harvested after four months was compared to analyze the differences in various morphological characteristics, biomass allocation, and physiological characteristics of the leaves of seedlings of J. regia f. luodianense, and a comprehensive evaluation of the plasticity indexes was conducted. The results showed that under moderate (50%) full light intensity, the leaf area, specific leaf area, leaf biomass, and chlorophyll content increased, and improved photosynthesis and promoted the accumulation of free proline content and peroxidase (POD) activity. The accumulation of malondialdehyde was also the lowest in this treatment, indicating that the plants had the strongest adaptability under this light intensity. Moreover, under high (75%) full light intensity, the above functional characteristics of plants showed good performance. Under low (25%) full light intensity, plants also had higher specific leaf area, leaf biomass, and photosynthetic parameters. However, under full light, the cell membrane permeability decreased, the chlorophyll accumulation was the lowest, and the photosynthetic index was seriously inhibited. Our results showed that the plasticity of morphological characters was greater than that of biomass allocation and physiological characters; POD activity and stomatal conductance were the highest, followed by leaf area and chlorophyll b, whereas the plasticity of palisade tissue/sponge tissue thickness and lower-epidermis thickness were the lowest. In summary, there are evident differences in the sensitivity and regulation mechanisms of morphological characteristics, biomass allocation, and physiological indices of the seedling leaves of J. regia f. luodianense in response to light intensity. During the stage of seedling establishment, only the plants in the bare ground under full light can be induced to show obvious inhibition of phenotypic traits. In contrast, the plants in the forest margins and gaps and under the forest habitats under light intensity can regulate their own characteristics to maintain their growth and development. The wide light range and strong plasticity of the species might be two of the important reasons for its existence in a highly heterogeneous karst habitat.
... Because excessive irradiance has a detrimental impact on photosynthetic tissues, plants must produce smaller and thicker leaves with higher leaf mass per area under high light conditions. This morphology allows heat dissipation, avoiding damage from overheating and high transpiration rates [35,36]. Conversely, shaded conditions result in increasing area and decreasing thickness of leaves [26,37], with low leaf mass per unit area [38]. ...
Background:
The non-structural carbohydrates (NSCs), carbon (C), nitrogen (N), and phosphorus (P) are important energy source or nutrients for all plant growth and metabolism. To persist in shaded understory, saplings have to maintain the dynamic balance of carbon and nutrients, such as leaf NSCs, C, N and P. To improve understanding of the nutrient utilization strategies between shade-tolerant and shade-intolerant species, we therefore compared the leaf NSCs, C, N, P in response to shade between seedlings of shade-tolerant Schima superba and shade-intolerant Cunninghamia lanceolate. Shading treatments were created with five levels (0, 40, 60, 85, 95% shading degree) to determine the effect of shade on leaf NSCs contents and C:N:P stoichiometry characteristics.
Results:
Mean leaf area was significantly larger under 60% shading degree for C. lanceolata while maximum mean leaf area was observed under 85% shading degree for S. superba seedlings, whereas leaf mass per area decreased consistently with increasing shading degree in both species. In general, both species showed decreasing NSC, soluble sugar and starch contents with increasing shading degree. However shade-tolerant S. superba seedlings exhibited higher NSC, soluble sugar and starch content than shade-intolerant C. lanceolate. The soluble sugar/starch ratio of C. lanceolate decreased with increasing shading degree, whereas that of S. superb remained stable. Leaf C:N ratio decreased while N:P ratio increased with increasing shading degree; leaf C:P ratio was highest in 60% shading degree for C. lanceolata and in 40% shading degree for S. superba.
Conclusion:
S. superba is better adapted to low light condition than C. lanceolata through enlarged leaf area and increased carbohydrate reserves that allow the plant to cope with low light stress. From mixed plantation viewpoint, it would be advisable to plant S. superba later once the canopy of C. lanceolata is well developed but allowing enough sunlight.
