Kranz Syndrome Research Articles

Atypical foliar anatomy related to Kranz syndrome in Paspalum inaequivalve and Paspalum microstachyum (Poaceae: Panicoideae: Paniceae)

Paspalum L. is a large and complex genus, enclosing more than 300 species, whose boundaries and infrageneric classification are still being studied. Recent phylogenetic analyses suggest that Paspalum inaequivalve Raddi and Paspalum microstachyum J. Presl, from the Inaequivalvia informal group, should be excluded from Paspalum. Focused on the unclear taxonomic position of P. inaequivalve and P. microstachyum, their leaf anatomy was studied, and some atypical features related to C 4 photosynthesis were found. This atypical Kranz syndrome is the aim of this research. Transverse leaf blade sections from fresh and herbarium material of P. inaequivalve were studied by light, fluorescence, and transmission microscopy. Additionally, sheaths and culms of P. inaequivalve and leaf blades of P. microstachyum were observed by light microscopy. δ 13C isotope discrimination was determinated for P. inaequivalve. We compared our results with available anatomical data from related taxa. As well as typical mesophyll cells (PCA) and mestome sheath cells (PCR), a third type of cells, here called ‘globose parenchymatous cells’, was observed in leaf blades of P. inaequivalve and P. microstachyum. These cells are placed externally to the mestome sheaths of the first and second vascular bundles, they have thin walls, with no developed suberine lamella, few chloroplasts with 1–2 starch grains, thylakoids not organized in grana, and a large central vacuole. The globose parenchymatous cells represent a novel trait in P. inaequivalve and P. microstachyum, further supporting the close relation between both species and their exclusion from the genus Paspalum. This atypical Kranz syndrome has not been described in Paspalum before, but the globose parenchymatous cells here described resemble the distinct cells considered as remnants of the outer parenchymatous sheath described for Anthaenantiopsis, some sections of Panicum L., and Chaetium Nees, providing possible taxonomic significance.

Screening of certain mangroves for photosynthetic carbon metabolic pathway

The mangroves Rhizophora lamarkii, Ceriops roxburghiana, Bruguiera gymnorrhiza, Aegiceras corniculatum, and Lumnitzera racemosa were screened for their carbon metabolic pathways by measuring net photosynthetic rate (P N), 13C discrimination rate, leaf anatomy, titratable acidity, and activities of phosphoenolpyruvate carboxylase, NADH-malate dehydrogenase, alanine aminotransferase, aspartate aminotransferase, and pyruvate phosphate dikinase. The tested mangroves had a well developed succulence, opening of stomata during day time and closure in the night hours, and absence of diurnal fluctuation of organic acids in their leaves which excludes the possibility of these species being CAM plants. Moreover, the leaf anatomy had not exhibited Kranz syndrome. The high values of discrimination against 13C, low P N, high CO2 compensation concentration, and the activities of aminotransferases in the direction of alanine formation suggest that the species may follow C3 mode of carbon metabolic pathway.

Anatomia e ultra-estrutura foliar de Cyperus maritimus Poir. (Cyperaceae): estratégias adaptativas ao ambiente de dunas litorâneas

Foram analisados aspectos anatômicos e ultra-estruturais foliares de Cyperus maritimus Poir. visando identificar caracteres adaptativos ao ambiente de dunas litorâneas. Para isto, indivíduos ocorrentes nas dunas da Praia de Pipa, Rio Grande do Norte, Brasil, tiveram as folhas submetidas a diferentes análises: microscopia de luz, eletrônica de varredura e de transmissão. Em vista frontal, a epiderme possui tricomas unicelulares, corpos de sílica, cutícula estriada e depósitos de cera. Em secção transversal, a epiderme é unisseriada com parede periclinal externa espessa. Os estômatos são paracíticos, podendo o ostíolo estar obstruído por cera. Abaixo da face adaxial observa-se o parênquima aqüífero. Os feixes vasculares ocorrem em múltiplas fileiras, sendo as periféricas compostas por feixes de diâmetro menor que os feixes da fileira central. Foram observados caracteres relacionados à síndrome Kranz do tipo clorociperóide. Aspectos ultra-estruturais, como cloroplastos presentes na bainha do feixe, esses sem grana evidentes, entre outras características, reforçam a possível ocorrência da via C4 nesta espécie. É a primeira citação para o gênero Cyperus de cloroplasto nas celulas do tecido vascular. Neste trabalho foi possível observar vários caracteres importantes para plantas submetidas a estresse hídrico e salino, como: cera, células epidérmicas com parede periclinal externa espessada, parênquima aqüífero e síndrome Kranz.

Classification and Biogeography of Panicoideae (Poaceae) in the New World

Panicoideae (Poaceae) in the New World comprise 107 genera (86 native) and 1357 species (1248 native). As circumscribed herein, Panicoideae include eight tribes: Andropogoneae, Arundinelleae, Centotheceae, Gynerieae, Isachneae, Paniceae, Steyermarkochloeae, and Thysanolaeneae. The two major tribes are Andropogoneae with 230 species (16.95% of all New World panicoids), and Paniceae with 1082 species (79.73%). Andropogoneae are divided into nine subtribes (Andropogoninae, Anthistiriinae, Coicinae, Germainiinae, Ischaeminae, Rottboelliinae, Saccharinae, Sorghinae, and Tripsacinae), while Paniceae are divided into seven subtribes (Arthropogoninae, Cenchrinae, Digitariinae, Melinidinae, Panicinae, Paspalinae, and Setariinae). Brazil is the center of diversity of New World panicoids with 741 species (54.6% of all species). The distributions of endemic and restricted taxa were analyzed in terms of biogeographical regions and Kranz syndrome. Amphitropical disjuncts and widespread species are also discussed.

Arquitetura, anatomia e histoquímica das folhas de Gomphrena arborescens L.f. (Amaranthaceae)

As folhas de Gomphrena arborescens L.f., nativa do Cerrado, são utilizadas contra dismenorréia na medicina popular. As investigações morfológica, anatômica e histoquímica das folhas visam fornecer elementos para taxonomia, para controle de qualidade de drogas e identificação microscópica de fitoterápicos. Folhas foram coletadas em Brasília, DF, nas áreas de Cerrado do Centro Olímpico da Univesidade de Brasília e na Reserva Ecológica do IBGE. Foram efetuadas análises morfológicas, anatômicas e histoquímicas. Os estudos de arquitetura foliar, densidade estomática e de vênulas terminais foram efetuados em folhas diafanizadas, apresentando médias de 42,39±15,50 estômatos mm-2 e 11,7±3,55 vênulas terminais mm². As folhas apresentaram duas formas de tricomas tectores. O mesofilo bilateral apresenta tecido clorenquimático disposto radialmente à bainha parenquimática completa que envolve os feixes vasculares, características da síndrome Kranz associada ao ciclo fotossintético C4. O amido concentra-se na bainha parenquimática. As folhas são ricas em celulose, drusas de oxalato de cálcio, proteínas, lignina e outros compostos fenólicos, entre eles os flavonóides. As folhas apresentam morfologia variável, mas o padrão anatômico e de venação são constantes.

Phylogenetics of Paniceae (Poaceae)

Paniceae demonstrate unique variability of photosynthetic physiology and anatomy, including both non-Kranz and Kranz species and all subtypes of the latter. This variability suggests hypotheses of independent origin or reversals (e.g., from C(4) to C(3)). These hypotheses can be tested by phylogenetic analysis of independent molecular characters. The molecular phylogeny of 57 species of Paniceae was explored using sequences from the grass-specific insert found in the plastid locus rpoC2. Phylogenetic analyses confirmed some long-recognized alliances in Paniceae, some recent molecular phylogenetic results, and suggested new relationships. Broadly, Paniceae were found to be paraphyletic with Andropogoneae, Panicum was found to be polyphyletic, and Oplismenus hirtellus was resolved as the sister group to the remaining ingroup species. A particularly well-supported clade in the rpoC2 tree included four genera with non-Kranz species and three with distinctively keeled paleas. As previously suggested, the PCK (phosphoenol pyruvate carboxykinase) C(4) subtype arose once within Paniceae. All clades with non-Kranz species had Kranz ancestors or sister taxa suggesting repeated loss of the Kranz syndrome.

Vegetational patterns within coastal salines in southern Yemen

AbstractFor the first time, the species diversity and zonation of coastal dunes and littoral salt marshes of the extensive coastal belt of southern Arabia is analyzed by means of two transects (Aden/Madinat as‐Shab; al‐Mukalla/Feleq). The vegetation analysis results five (Aeluropus lagopoides‐, Halopeplis perfoliata‐, Halopyrum mucronatum‐, Sevada schimperi‐association, and one rest group – Aden transect) respectively seven (Anabasis ehrenbergii‐, Arthrocnemum macrostachyum‐, Cyperus conglomeratus‐, Stipagrostis paradisea‐, Urochondra setulosa‐Arthrocnemum macrostachyum‐, Urochondra setulosa‐association, and one rest group – Feleq transect) homogenous ecological groups, which form clearly defined belts following a strong edaphical gradient. While the associations in the vicinity of Aden are in good accordance with those, reported from the coasts of the Red Sea and the Arabian Gulf, three new associations, partly rich in endemics are reported from the southern coast of Hadramout (Feleq transect). Due to the plantgeographical relationships of the taxa (Eritreo‐Arabian chorotype), they are strongly related to the coasts of NE Africa. In addition, Coelachyrum piercei is reported as new to the flora of Yemen, closing herewith a gap in the distribution pattern of this Omano‐Makranian taxon.All taxa were examined with respect to their photosynthetic pathway type (Kranz syndrome, δ13C‐values) to clarify, if there is a correlation between the distribution of C3‐/C4‐plants and the ecological requirements of the halophytes. It turned out, that hygrohalophytes and oligohaline, xerix associations are dominated by C3‐plants, whereas C4‐plants are typical of xerohalophytic associations and those, showing an intermediate position with respect to soil salinity and available water.

Anatomy of the Muhlenbergia montana Complex (Poaceae)

Leaf anatomical data from 13 taxa of the Muhlenbergia montana complex were surveyed, and 18 critical characters were found. Results obtained from the phenetic analysis of anatomical data have given independent data to compare with the morphological, phytogeographical, and flavonoid data (previously reported) in determining the relationships of the species. An anatomical dendogram illustrates a close corroboration. Leaf anatomy clearly supports the recognition of a close relationship among the two taxa-M. virescens and M. quadridentata-as has been suggested in previous reports. Therefore, a change of level is suggested in which M. quadridentata (H.B.K.) Kunth is proposed here as M. virescens (H.B.K.) Kunth ssp. quadridentata (H.B.K.) Herrera (comb. nov.). Approximately 160 species of Muhlenbergia Schreb. are reported by Clayton and Renvoize (1986). The majority of the species are xeromorphic, caespitose forms that grow in steppe or desert climates of the western United States, Mexico, and Central and South America (Pohl, 1969). Members of the genus occur from subarctic North America to southern South America (Pohl, 1969) and show diversity in growth habit, morphology, and ecology. The Kranz syndrome C4 type leaf anatomy is reported *for the genus (Hattersley, 1984), particularly the parenchyma sheath subtype which is the most common and which is found in species occupying the most arid regions (Brown, 1977).

Infrageneric phenetic relations in new world Panicum (Poaceae: Panicoideae: Paniceae): a numerical analysis

Numerical studies based on 64 characters placed 188 American species of the genus Panicum L. in 23 sections clustered in five major groups. The same major groups were obtained when the five characters that define the subgenera were excluded from the analyses. Groups 1 and 2 included C4 species. Species with an outer parenchymatous sheath in group 1 corresponded to subg. Panicum, whereas species with a single mestome sheath in group 2 corresponded to subg. Agrostoides. Group 3 included the intermediate C3–C4 species belonging to subg. Steinchisma. C3 sections were clustered together but arranged in a different way than previous classifications. Group 4 included sections from subg. Phanopyrum with basic chromosome number x = 10, whereas group 5 included the other sections of Phanopyrum with x = 9 together with sections from subg. Dichanthelium also with x = 9. The two subgroups of subg. Phanopyrum are not only different in chromosome number but also in inflorescence type and upper anthecium ornamentation. On the other hand the x = 9 sections of Phanopyrum remain in the same cluster with sections from subg. Dichanthelium in numerical analyses where basic chromosome number is removed. These results suggest the possibility of sinking Dichanthelium into Phanopyrum and then split the latter when more cytological data become available. Kranz syndrome and basic chromosome number were the characters with the highest contribution to the delimitation of the major groups. On the basis of these results different existing hypotheses on the classification of the genus Panicum are discussed. Key words: Panicum, numerical taxonomy.

Photosynthetic Pathways and Ecological Distribution of Halophytes from Four Littoral Salt Marshes (Egypt/Sinai, Saudi Arabia, Oman and Iran)

Summary This is the first comparative study to be made of four littoral salt marshes in South-West Asia (Shura Arwashie, Egypt/Sinai; Tawwal, Saudi Arabia; Qurm, Oman; Bandar-e Khamer, Iran). The aim was to find out whether there is a relation between the photosynthetic pathways and the ecological distribution of the halophytes. To answer this question, quantitative-analytic survey techniques were used to determine the zonation of the littoral salt marshes, and the species making up the vegetation zones were examined with regard to their photosynthetic pathways (Kranz syndrome, δ13C values). Attention was given to soil moisture and soil salinity. With the aid of photosynthesis spectra the proportional occurrence of each group in the zones could be determined. It turned out that in all four examples there is a correlation between photosynthetic pathway, soil moisture and soil salinity. The C4 plant distribution is not only related to high summer and winter temperatures and to soil moisture, but is also influenced by salinity. The C4 species colonize a transition zone between C3 hygrohalophytes and xerophytic, nonhalophytic C3 species.

The Kranz syndrome in the Eragrostideae (Chloridoideae, Poaceae) as indicated by carbon isotopic ratios*

13C/12C ratios are generally regarded as being very reliable indicators of C3 or C4 photosynthesis. These relative carbon isotope ratios are expressed as a negative δ 3C and fall into two distinct groups: Kranz (or C4) plants with δ between -9°/00 no and -18°/00 and non-Kranz (C3) plants with δ between -22°/00 and -280/00 no. In this paper, 29 taxa, representing 12 genera, of the tribe Eragrostideae were examined by mass spectrometry for their δ 13C in dried leaf tissue. All these taxa proved to be C4, plants with δ13C values ranging between -13,6°/oo and -10.9°/oo. These findings confirmed published leaf anatomical observations which showed that all the studied taxa had characteristic Kranz leaf anatomy.

A C4 plant from the Pliocene

THE Kranz syndrome1,2 is a group of physiological and anatomical features occurring in certain angiosperms; the grasses in particular show increasing efficiency in CO2 assimilation. Recent research3,4 suggests an origin of the syndrome in South America and its subsequent dominance in other continents. The Kranz syndrome occurs in all plants fixing CO2 by phosphoenolpyruvate carboxylase in the mesophyll and subsequent production of Calvin–Benson cycle intermediates in the bundle sheath5. The physiological processes in C4 plants are closely related to their specialised leaf anatomy known as Kranz anatomy6 by which they are distinguished from C3 plants. It is suggested that ancestral non-Kranz C3 grasses originated during the Cretaceous period4. We report here fossil evidence indicating that C4 plants occurred at least during the late Tertiary period.

DISTRIBUTION OF KRANZ SYNDROME AMONG ASTERACEAE

A broad survey of the Asteraceae, involving visual characters, was undertaken to ascertain the occurrence of possible Kranz‐type (C4) plants. One hundred and seven taxa representing 51 genera were examined by mass spectrometry for their 13C/12C ratios in dried leaf tissue. Species possessing the high 13C/12C ratio typical of the Kranz syndrome were found in genera belonging to the following tribes: Heliantheae (subtribe Coreopsideae)—Chrysanthellum (4 spp.), Eryngiophyllum (1 sp.), Glossocardia (1 sp.), Glossogyne (1 sp.), Isostigma (5 spp.); Tageteae—Pectis (21 spp.); and Helenieae—Flaveria (4 spp.). The only genus to possess both C3 and C4 species was Flaveria in which all but four species appear to possess a C3‐type metabolism. When appropriate, the phyletic input of these data is discussed.

Distribution of Kranz Syndrome Among Asteraceae

A broad survey of the Asteraceae, involving visual characters, was undertaken to ascertain the occurrence of possible Kranz-type (C4) plants. One hundred and seven taxa representing 51 genera were examined by mass spectrometry for their 13C/12C ratios in dried leaf tissue. Species possessing the high 13C/12C ratio typical of the Kranz syndrome were found in genera belonging to the following tribes: Heliantheae (subtribe Coreopsideae)—Chrysanthellum (4 spp.), Eryngiophyllum (1 sp.), Glossocardia (1 sp.), Glossogyne (1 sp.), Isostigma (5 spp.); Tageteae—Pectis (21 spp.); and Helenieae—Flaveria (4 spp.). The only genus to possess both C3 and C4 species was Flaveria in which all but four species appear to possess a C3-type metabolism. When appropriate, the phyletic input of these data is discussed.

SYSTEMATICS OF PHOTOSYNTHETIC CARBON FIXATION PATHWAYS IN EUPHORBIA

SummaryUsing the technique of carbon isotopic ratio determinations, 20 selected species of Euphorbia (s. lat.) have been investigated to determine their metabolic status. Previous work is reviewed, and as found by previous investigators, enriched 13C values ‐ indicating Crassulacean acid metabolism or C4 photosynthesis ‐ have been detected only in subgenera Euphorbia and Chamaesyce, respectively. All taxa investigated in subg. Chamaesyce except two species of subsect. Acutae show the Kranz syndrome and C4 photosynthesis; the probable phylogenetic origin of the subgenus is therefore pinpointed within this subsection. Apparently taxa with Crassulacean acid metabolism and C4 photosynthesis have arisen independently within the genus.

The Kranz Syndrome in the Gramineae as Indicated by Carbon Isotopic Ratios

The Kranz syndrome, as indicated by relatively high 13C/12C ratios is characteristic of 16 ½ tribes and about ½ of the species of the Gramineae. Data are given for 198 species from 129 genera and 47 tribes, and from at least 6 subfamilies of grasses. This information is correlated with data from the literature on anatomical and physiological characters of both Kranz and non-Kranz grasses. All subfamilies, tribes, and genera seem to be uniformly all Kranz or non-Kranz except the subfamily Panicoideae and the genus Panicum which have both Kranz and non-Kranz species represented.

THE KRANZ SYNDROME IN THE GRAMINEAE AS INDICATED BY CARBON ISOTOPIC RATIOS

The Kranz syndrome, as indicated by relatively high 13C/12C ratios is characteristic of 16 ½ tribes and about ½ of the species of the Gramineae. Data are given for 198 species from 129 genera and 47 tribes, and from at least 6 subfamilies of grasses. This information is correlated with data from the literature on anatomical and physiological characters of both Kranz and non‐Kranz grasses. All subfamilies, tribes, and genera seem to be uniformly all Kranz or non‐Kranz except the subfamily Panicoideae and the genus Panicum which have both Kranz and non‐Kranz species represented.

Grass evolution, the Kranz syndrome, 13C/12C ratios, and continental drift.

THE Kranz syndrome is a group of related anatomical, cytological and physiological characters reported one way or another in about 10 families of angiospermous plants. The syndrome centres around the CO2 fixation aspect of photosynthesis, including synthesis of C4-dicarboxylic acids as the initial products of CO2 assimilation, utilization of most of the available CO2 producing a high 13C/12C ratio, having maximum efficiency at high temperatures, and all related to a sheath around the bundles in the leaf. All present evidence indicates that detection of any one of the aspects of the syndrome denotes the whole syndrome1. The syndrome is uncommon among plants including angiosperms and is therefore considered to be the evolved or derived condition. Although in Atriplex2 it seems to have evolved recently within the genus, in the Gramineae the evolution of the Kranz syndrome seems to have occurred only two or three times early in grass evolution. This assumption, which is fundamental in this discussion, is based on the uniformity of tribes and subfamilies as Kranz or non-Kranz, and the proven predictability of the syndrome from single aspects of it (in this work from the 13C/12C ratio), from various other characters, or from classification3. The only exception reported, of Kranz and non-Kranz species within one grass genus, is within Panicum4 where the subgenus Dichanthelium is non-Kranz whereas the more typical species of the subgenus Eupanicum are characterized by the Kranz syndrome (Table 1). Recently we found, by determination of 13C/12C ratios of over 180 species, in 110 genera, of forty tribes of grasses, evidence of both types also among species of Eupanicum as well as among genera of the tribe Paniceae and among tribes of the subfamily Panicoideae (Table 1). It is this diversity of panicoid tribes (many of which are small and endemic), of the tribe Paniceae, and the genus Panicum that provides clues to the likely evolution of the grass family, superimposed, of course, on a great deal of other morphological, anatomical, cytological, and physiological evidence3,5. Furthermore, now that geology has accepted Wegenerian drifting continents and the break-up of an ancient single world-continent (Pangaea) by rifting under the concept of plate tectonics, evolution of the Gramineae can be discussed in terms of that concept; that is, Upper Jurassic–Middle Cretaceous separation of East Gondwana and the Middle to Late Cretaceous separation of South America from Africa, with the tropical belt approximately in its present relationship to the continents since Early Jurassic time6.

A Study of Some Anatomical Characters of North American Gramineæ. I

Previous articleNext article FreeA Study of Some Anatomical Characters of North American Gramineæ. ITheo. HolmTheo. Holm Search for more articles by this author PDFPDF PLUS Add to favoritesDownload CitationTrack CitationsPermissionsReprints Share onFacebookTwitterLinkedInRedditEmail SectionsMoreDetailsFiguresReferencesCited by Volume 16, Number 6Jun., 1891 Article DOIhttps://doi.org/10.1086/326666 Views: 51 Citations: 4Citations are reported from Crossref Journal History This article was published in the Botanical Gazette (1876-1991), which is continued by International Journal of Plant Sciences (1992-present). PDF download Crossref reports the following articles citing this article:Marius-Nicușor Grigore, Constantin Toma Bulliform Cells, (Oct 2017): 325–338.https://doi.org/10.1007/978-3-319-66480-4_8P. R. Espie, H. E. Connor, I. J. McCracken Leaf-blade crimping in grasses: A new measure of growth, Experientia 48, no.11 (Jan 1992): 91–94.https://doi.org/10.1007/BF01923616Bruce N. Smith, Walter V. Brown THE KRANZ SYNDROME IN THE GRAMINEAE AS INDICATED BY CARBON ISOTOPIC RATIOS, American Journal of Botany 60, no.66 (Jul 1973): 505–513.https://doi.org/10.1002/j.1537-2197.1973.tb05951.xDietrich Brandis V. Remarks on the Structure of Bamboo Leaves., Transactions of the Linnean Society of London. 2nd Series: Botany 7, no.55 (Apr 2009): 69–92.https://doi.org/10.1111/j.1095-8339.1907.tb00152.x