A síndrome Kranz é um conjunto de características anatômicas, ultra-estruturais e bioquímicas que culminam na realização da fotossíntese C4. Tal síndrome apresenta grande diversidade dentre as Angiospermas, tornando-se conveniente seu estudo em todos os níveis acima citados para a completa caracterização da mesma. No presente trabalho foi investigada a síndrome Kranz de Gomphrena arborescens e G. scapigera (Amaranthaceae) com ênfase na origem ontogenética da bainha Kranz, na descrição ultra-estrutural e na confirmação bioquímica sobre o tipo de fotossíntese C4. O desenvolvimento foliar dessas espécies indica que a bainha Kranz é originada da camada mais interna do mesofilo, a endoderme foliar. Uma discussão sobre os termos presentes na literatura para a descrição dessa bainha, todos eles focados em sua função na fotossíntese C4, demonstra a importância de se utilizar termos que informem a origem ontogenética dessa bainha, como endoderme e periciclo. Na análise ultra-estrutural, foram identificados possíveis fatores que interferem na fotossíntese de ambas as espécies, como o espessamento e a composição da parede da bainha Kranz, o posicionamento centrípeto dos cloroplastos e a presença de retículo periférico nos mesmos. Embora a análise bioquímica tenha resultado em informações ainda não conclusivas, o dimorfismo dos cloroplastos sugere a realização da fotossíntese C4 do tipo NADP-ME. O presente trabalho, de uma forma geral, contribui ao conhecimento da síndrome Kranz dentre as Amaranthaceae s.s., um grupo em que a ultra-estrutura e a bioquímica ainda são pouco conhecidas, e ressalta a importância dos estudos anatômicos, principalmente com enfoque ontogenético, para o melhor conhecimento da diversidade da síndrome Kranz dentre as Angiospermas. / The Kranz syndrome is a set of anatomical, ultrastructural and biochemical features that culminate in the C4 photosynthesis. This syndrome has a huge diversity among Angiosperms, so it became suitable to survey all the levels above cited for its complete characterization. In the present work the Kranz syndrome of Gomphrena arborescens and G. scapigera (Amaranthaceae) is studied, with emphasis on the ontogenetic origin of the Kranz sheath, on the ultrastructural description, and on the biochemical confirmation about the C4 photosynthesis kind. The foliar development of these species shows that the Kranz sheath is originated from the inner layer of the mesophyll, the foliar endodermis. A discussion about the literature terms used to describe the Kranz sheath, all of them referring to the function of this layer in C4 photosynthesis, demonstrates the importance of using terms that inform the ontogenetic origin of this layer, such as endodermis and perycicle. The ultrastructural analysis identified possible factors that interfere on the C4 photosynthesis of both species, such as wall thickening and composition of Kranz sheath cells, the centripetal position of chloroplasts and the peripheral reticulum in chloroplasts. Although biochemical analysis has resulted in no conclusive information, the chloroplast dimorphism suggests the NADP-ME C4 photosynthesis. This work, in a general way, contributes to the knowledge of the Kranz syndrome among Amaranthaceae s.s., a group that has the ultrastructure and the biochemistry of C4 photosynthesis poorly known. It also draws attention to the importance of anatomical surveys concerning the ontogenetic origin of Kranz sheath for a better understanding on the diversity of Kranz syndrome among Angiosperms.

A síndrome Kranz é um conjunto de características anatômicas, ultra-estruturais e bioquímicas que culminam na realização da fotossíntese C4. Tal síndrome apresenta grande diversidade dentre as Angiospermas, tornando-se conveniente seu estudo em todos os níveis acima citados para a completa caracterização da mesma. No presente trabalho foi investigada a síndrome Kranz de Gomphrena arborescens e G. scapigera (Amaranthaceae) com ênfase na origem ontogenética da bainha Kranz, na descrição ultra-estrutural e na confirmação bioquímica sobre o tipo de fotossíntese C4. O desenvolvimento foliar dessas espécies indica que a bainha Kranz é originada da camada mais interna do mesofilo, a endoderme foliar. Uma discussão sobre os termos presentes na literatura para a descrição dessa bainha, todos eles focados em sua função na fotossíntese C4, demonstra a importância de se utilizar termos que informem a origem ontogenética dessa bainha, como endoderme e periciclo. Na análise ultra-estrutural, foram identificados possíveis fatores que interferem na fotossíntese de ambas as espécies, como o espessamento e a composição da parede da bainha Kranz, o posicionamento centrípeto dos cloroplastos e a presença de retículo periférico nos mesmos. Embora a análise bioquímica tenha resultado em informações ainda não conclusivas, o dimorfismo dos cloroplastos sugere a realização da fotossíntese C4 do tipo NADP-ME. O presente trabalho, de uma forma geral, contribui ao conhecimento da síndrome Kranz dentre as Amaranthaceae s.s., um grupo em que a ultra-estrutura e a bioquímica ainda são pouco conhecidas, e ressalta a importância dos estudos anatômicos, principalmente com enfoque ontogenético, para o melhor conhecimento da diversidade da síndrome Kranz dentre as Angiospermas. / The Kranz syndrome is a set of anatomical, ultrastructural and biochemical features that culminate in the C4 photosynthesis. This syndrome has a huge diversity among Angiosperms, so it became suitable to survey all the levels above cited for its complete characterization. In the present work the Kranz syndrome of Gomphrena arborescens and G. scapigera (Amaranthaceae) is studied, with emphasis on the ontogenetic origin of the Kranz sheath, on the ultrastructural description, and on the biochemical confirmation about the C4 photosynthesis kind. The foliar development of these species shows that the Kranz sheath is originated from the inner layer of the mesophyll, the foliar endodermis. A discussion about the literature terms used to describe the Kranz sheath, all of them referring to the function of this layer in C4 photosynthesis, demonstrates the importance of using terms that inform the ontogenetic origin of this layer, such as endodermis and perycicle. The ultrastructural analysis identified possible factors that interfere on the C4 photosynthesis of both species, such as wall thickening and composition of Kranz sheath cells, the centripetal position of chloroplasts and the peripheral reticulum in chloroplasts. Although biochemical analysis has resulted in no conclusive information, the chloroplast dimorphism suggests the NADP-ME C4 photosynthesis. This work, in a general way, contributes to the knowledge of the Kranz syndrome among Amaranthaceae s.s., a group that has the ultrastructure and the biochemistry of C4 photosynthesis poorly known. It also draws attention to the importance of anatomical surveys concerning the ontogenetic origin of Kranz sheath for a better understanding on the diversity of Kranz syndrome among Angiosperms.

C4 plants have a biochemical carbon concentrating mechanism (CCM) that increases CO2 concentration around Rubisco in the bundle sheath (BS). Maize CCM has two CO2 delivery pathways to the Bundle Sheath (BS) (respectively via malate, MAL or aspartate, ASP); rates of PGA reduction, carbohydrate synthesis and PEP regeneration vary between BS and Mesophyll (M) cells. For these anatomical and biochemical complexities, C4 plants are highly sensitive to light conditions. Under limiting light, the activity of the CCM generally decreases, causing an increase in leakiness, (Φ), the ratio of CO2 retrodiffusing from the BS relative to C4 carboxylation processes. This increase in Φ had been theoretically associated with a decrease in biochemical operating efficiency (expressed as ATP cost of gross assimilation, ATP / GA) under low light and, because a proportion of canopy photosynthesis is carried out by shaded leaves, to potential productivity losses at field scale. In C4 leaves, because of the concentric anatomy, light reaches M cells before the deeper BS (Evans et al., 2007), and could alter the energetic partitioning balance between BS and M and potentially cause efficiency losses. In this experimental programme I investigated strategies deployed by C4 plants to adjust operating efficiency under different illumination conditions. Firstly, maize plants were grown under high and low light regimes (respectively HL, 600 vs LL, 100 μE m-2 s-1). Short term acclimation of Φ was compared from isotopic discrimination (Δ), gas exchange and photochemistry using an improved modelling approach which does not suffer from elements of circularity. Long term acclimation to low light intensities brought about physiological changes which could potentially increase the operating efficiency under limiting ATP supplies. Secondly, profiles of light penetration across a leaf were used to derive the potential ATP supply for M and BS cells induced by changing light quality. Empirical measurements of net CO2 uptake, ATP production rate and carbon isotope discrimination were made on plants under a low light intensity. The overall conversion efficiency was not affected by light quality. A comprehensive metabolic model highlighted the importance of both CO2 delivery pathways in maize. Further, metabolic plasticity allowed the balancing of ATP and NADPH requirements between BS and M. Finally, I tested the hypothesis that plants can modify their physiology so as to reach a status of higher operating efficiency when exposed to high light and then to low light, so as to mimic the transition which leaves undergo when shaded by newly emerging leaves in a crop canopy. Plants were grown under high light and low light for three weeks, then, HL plants were transferred to low light for a further three weeks. Re-acclimation was very effective in reducing ATP cost of net assimilation under low light intensities. In addition, the hyperbolic leakiness increase observed under low light intensities was not associated to operating efficiency loss. Overall, in the three experimental Chapters I showed compelling theoretical and empirical evidence proving the hypothesis that C4 plants deal with low light conditions and with different light qualities without losing operating efficiency.

Orientador: Tatiane Maria Rodrigues / Resumo: Em folhas de diversas espécies vegetais, a bainha de células que envolve os feixes vasculares se estende em direção à epiderme em ambas as faces do limbo formando extensões da bainha do feixe (EBF), o que caracteriza folhas heterobáricas. Folhas que não apresentam EBF são chamadas homobáricas. Pesquisas demostram que folhas homobáricas garantem maior difusão lateral de gases, enquanto as EBF podem atuar na condução de água e aumentar o desempenho fotossintético foliar garantindo a transferência de luz para as camadas mais internas do mesofilo. As diferenças morfofuncionais entre os dois tipos foliares parecem estar relacionadas ao ambiente onde as plantas ocorrem e à sua distribuição nos diferentes estratos vegetacionais. Entretanto, para formações vegetais no Brasil não foram encontrados estudos sobre a distribuição de espécies com folhas homobáricas e heterobáricas. Esse trabalho teve como objetivo estudar a anatomia foliar de espécies herbáceas, arbustivas e arbóreas na floresta estacional semidecídua, cerradão e cerrado stricto sensu a fim de identificar folhas homobáricas e heterobáricas, buscando associar a presença/ausência de EBF com o porte das plantas e com o ambiente. Quadrantes de 20mX20m foram demarcados nas fragmentos vegetacionais. Folhas completamente expandidas foram coletadas de 59 espécies no cerradão (32 arbóreas; 19 arbustos e 8 herbáceas); 68 espécies no cerrado stricto sensu (21 arbóreas; 30 arbustos e 17 herbáceas) e 51 espécies na floresta estacional ... (Resumo completo, clicar acesso eletrônico abaixo) / Abstract: In leaves of several plants species, the vascular bundle sheath extends toward the epidermis on both sides of the blade forming bundle sheath extensions (BSE), characterizing heterobarics leaves. Leaves lacking BSE are called homobarics. Research shows that homobaric leaves provide greater lateral movements of gases, while the BSE can act conducting water and increasing the leaf photosynthetic performance, what ensures the transference of light to the inner layers of the mesophyll. The morphological and functional differences between the two leaf types seem to be related to the environment where the plants occur and their distribution in the different vegetation strata. However, there are no studies on the distribution of species with homobarics and heterobarics leaves for vegetation formations in Brazil. This work aimed to study the leaf anatomy of herbs, shrubs and trees in semideciduous seasonal forest, cerradão and cerrado stricto sensu to identify homobarics and heterobarics leaves, searching to associate the presence / absence of BSE to the plant strata and environmental characteristics. Quadrants of 20mX20m were marked in vegetation fragments. Fully expanded leaves were collected from 59 species in the cerradão (32 tree, 19 shrubs and 8 herbs); 68 species in the cerrado stricto sensu (21 tree, 30 shrubs and 17 herbs) and 51 species in the semideciduous seasonal forest (23 tree, 17 shrubs and 11 herbs). Samples of the leaf blade were processed according to standard tech... (Complete abstract click electronic access below) / Mestre

In this Ph.D. thesis I am dealing with the effect of dynamic irradiance on (i) photosynthesis, growth and bundle sheath leakiness (for CO2) of C4 plants and (ii) bryophyte photosynthesis. Part of this thesis is literature review on broader aspects of the dynamic light effects on photosynthetic and growth processes in plants.