New insights into the role of the root system of epiphytic bromeliads: comparison of root and leaf trichome functions in acquisition of water and nutrients.

BACKGROUND: In epiphytic bromeliads, the roots used to be considered poorly functional organs in the processes of absorption and metabolization of water and nutrients, while the leaves always acted as protagonists in both functions. More recent discoveries have been changing this old view of the root system. SCOPE: In this review, we will address the old thoughts of the scientific community regarding the function performed by the roots of epiphytic bromeliads (mere holdfast structures with low physiological activity) and the importance of a reduced or lack of root system for the emergence of epiphytism. We will present indirect and direct evidence that contradicts this older hypothesis. Furthermore, the importance of the root absorptive function mainly for juvenile tankless epiphytic bromeliads and the characteristics of the root absorption process of adult epiphytic tank bromeliads will be thoroughly discussed in physiological aspects. Finally, some factors (species, substrate, environmental conditions) that influence the absorptive capability of the roots of epiphytic tank bromeliads will also be considered in this review, highlighting the importance that the absorptive role of the roots have for the plasticity of bromeliads that live on trees, which is an environment characterized by the intermittent availability of water and nutrients. CONCLUSIONS: The roots of tank-forming epiphytic bromeliads play important roles in the absorption and metabolization of nutrients and water. The importance of roots stands out mainly for juvenile tankless bromeliads since the root is the main absorptive organ. In larger plants with tank, although the leaves become the protagonists in the resource acquisition process, the roots complement the absorptive function of the leaf trichomes, resulting in a better growth of the bromeliad. The physiological and biochemical properties of the processes of absorption and distribution of resources in the tissues seem to differ between absorption by trichomes and roots.

Spatial patterns of photosynthesis in thin- and thick-leaved epiphytic orchids: unravelling C3-CAM plasticity in an organ-compartmented way.

BACKGROUND AND AIMS: A positive correlation between tissue thickness and crassulacean acid metabolism (CAM) expression has been frequently suggested. Therefore, this study addressed the question of whether water availability modulates photosynthetic plasticity in different organs of two epiphytic orchids with distinct leaf thickness. METHODS: Tissue morphology and photosynthetic mode (C3 and/or CAM) were examined in leaves, pseudobulbs and roots of a thick-leaved (Cattleya walkeriana) and a thin-leaved (Oncidium 'Aloha') epiphytic orchid. Morphological features were studied comparing the drought-induced physiological responses observed in each organ after 30 d of either drought or well-watered treatments. KEY RESULTS: Cattleya walkeriana, which is considered a constitutive CAM orchid, displayed a clear drought-induced up-regulation of CAM in its thick leaves but not in its non-leaf organs (pseudobulbs and roots). The set of morphological traits of Cattleya leaves suggested the drought-inducible CAM up-regulation as a possible mechanism of increasing water-use efficiency and carbon economy. Conversely, although belonging to an orchid genus classically considered as performing C3 photosynthesis, Oncidium 'Aloha' under drought seemed to express facultative CAM in its roots and pseudobulbs but not in its leaves, indicating that such photosynthetic responses might compensate for the lack of capacity to perform CAM in its thin leaves. Morphological features of Oncidium leaves also indicated lower efficiency in preventing water and CO2 losses, while aerenchyma ducts connecting pseudobulbs and leaves suggested a compartmentalized mechanism of nighttime carboxylation via phosphoenolpyruvate carboxylase (PEPC) (pseudobulbs) and daytime carboxylation via Rubisco (leaves) in drought-exposed Oncidium plants. CONCLUSIONS: Water availability modulated CAM expression in an organ-compartmented manner in both orchids studied. As distinct regions of the same orchid could perform different photosynthetic pathways and variable degrees of CAM expression depending on the water availability, more attention should be addressed to this in future studies concerning the abundance of CAM plants.

Nitrogen metabolism in leaves of a tank epiphytic bromeliad: characterization of a spatial and functional division.

The leaf is considered the most important vegetative organ of tank epiphytic bromeliads due to its ability to absorb and assimilate nutrients. However, little is known about the physiological characteristics of nutrient uptake and assimilation. In order to better understand the mechanisms utilized by some tank epiphytic bromeliads to optimize the nitrogen acquisition and assimilation, a study was proposed to verify the existence of a differential capacity to assimilate nitrogen in different leaf portions. The experiments were conducted using young plants of Vriesea gigantea. A nutrient solution containing NO3⁻/NH4⁺ or urea as the sole nitrogen source was supplied to the tank of these plants and the activities of urease, nitrate reductase (NR), glutamine synthetase (GS) and glutamate dehydrogenase (NADH-GDH) were quantified in apical and basal leaf portions after 1, 3, 6, 9, 12, 24 and 48 h. The endogenous ammonium and urea contents were also analyzed. Independent of the nitrogen sources utilized, NR and urease activities were higher in the basal portions of leaves in all the period analyzed. On the contrary, GS and GDH activities were higher in apical part. It was also observed that the endogenous ammonium and urea had the highest contents detected in the basal region. These results suggest that the basal portion was preferentially involved in nitrate reduction and urea hydrolysis, while the apical region could be the main area responsible for ammonium assimilation through the action of GS and GDH activities. Moreover, it was possible to infer that ammonium may be transported from the base, to the apex of the leaves. In conclusion, it was suggested that a spatial and functional division in nitrogen absorption and NH4⁺ assimilation between basal and apical leaf areas exists, ensuring that the majority of nitrogen available inside the tank is quickly used by bromeliad's leaves.

Specific leaf areas of the tank bromeliad Guzmania monostachia perform distinct functions in response to water shortage.

Leaves comprise most of the vegetative body of tank bromeliads and are usually subjected to strong longitudinal gradients. For instance, while the leaf base is in contact with the water accumulated in the tank, the more light-exposed middle and upper leaf sections have no direct access to this water reservoir. Therefore, the present study attempted to investigate whether different leaf portions of Guzmania monostachia, a tank-forming C(3)-CAM bromeliad, play distinct physiological roles in response to water shortage, which is a major abiotic constraint in the epiphytic habitat. Internal and external morphological features, relative water content, pigment composition and the degree of CAM expression were evaluated in basal, middle and apical leaf portions in order to allow the establishment of correlations between the structure and the functional importance of each leaf region. Results indicated that besides marked structural differences, a high level of functional specialization is also present along the leaves of this bromeliad. When the tank water was depleted, the abundant hydrenchyma of basal leaf portions was the main reservoir for maintaining a stable water status in the photosynthetic tissues of the apical region. In contrast, the CAM pathway was intensified specifically in the upper leaf section, which is in agreement with the presence of features more suitable for the occurrence of photosynthesis at this portion. Gas exchange data indicated that internal recycling of respiratory CO(2) accounted for virtually all nighttime acid accumulation, characterizing a typical CAM-idling pathway in the drought-exposed plants. Altogether, these data reveal a remarkable physiological complexity along the leaves of G. monostachia, which might be a key adaptation to the intermittent water supply of the epiphytic niche.

Ontogenic expression of TGFbeta 1, 2, and 3 and its receptors in the rat gastric mucosa.

The stomach of the rat undergoes extensive changes during the formation and maturation of gastric glands. The presence of transforming growth factor beta (TGFbeta) in rat milk and in the gastrointestinal tract of pups may suggest its role in this process. The current study evaluated the in vivo dynamic expression and distribution of TGFbeta1, beta2, beta3 and their receptors TbetaRI and TbetaRII in the gastric epithelium of 20-day fetal rats and 1-, 14-, 21-, and 30-day-old pups. Immunohistochemistry was used to detect the proteins, and staining was classified according to intensity and cell type. The results showed that the gastric epithelium expresses TGFbeta isoforms and receptors throughout development. We found that immunoreactivity paralleled the appearance of differentiated cells, such that surface mucous cells were the first to be immunostained and chief cells were the last. The intensity of reactions followed this same pattern, showing that the expression of TGFbeta isoforms spread along the gland with growth. Of interest, the highest apparent activity of TGFbeta was observed from 21 days onward, a period that is concomitant with weaning and maturation of most gastric cell types. In addition, surface mucous cells were strongly labeled at the basal cytoplasm at 14 days, suggesting an interaction with the connective tissue. In conclusion, the dynamic expression of TGFbeta1, beta2, beta3, and TbetaRI and TbetaRII through stomach development suggests significant paracrine and autocrine roles for this growth factor. We propose that temporal and spatial differences may be regulated by dietary changes, which in turn control cell proliferation and differentiation in the gastric epithelium.