Seasonal Cambial Activity and Formation of Secondary Phloem and Xylem in White Oaks (Quercus alba L.)

(1) Background: the cambium has seasonal activity, forming earlywood and early phloem with relatively wide conducting cells, which will function during the most favorable season, and latewood and late phloem with narrower conducting cells, which typically function during the less favorable season. However, few studies have focused on when these two contrasting tissue types are formed in relation to climatic conditions. (2) Methods: the senior author of this paper made weekly collections for an entire year of four specimens per collection back in the 1960s, using traditional anatomical methods to study in detail what the cambium was producing progressively. (3) Results: annual growth rings are evident in both secondary xylem and secondary phloem. The cambium resumes activity in early April, with simultaneous formation of wood and secondary phloem. Both latewood and late phloem production are initiated in early June, the peak of the favorable season. The cambium ends its activity in early August. Phloem growth rings are marked by radially narrow sieve elements interspersed among a band of axial parenchyma with dark contents. Most specimens produce only one fiber band per season. This feature may be used as an indirect phloem growth ring marker. Wood growth rings are marked by very wide vessels and thick-walled, radially narrow fibers. (4) Conclusions: growth rings are evident in both secondary xylem and secondary phloem. The trees produce their latewood and late phloem long before the beginning of autumn, indicating that they prepare ahead of the selective regime, a phenomenon most likely dependent on the photoperiod. Living sieve elements are present yearlong.

[1]  M. Olson,et al.  The vulnerability to drought-induced embolism-conduit diameter link: breaching the anatomy-physiology divide , 2023, IAWA Journal.

[2]  Kasia Ziemińska The role of imperforate tracheary elements and narrow vessels in wood capacitance of angiosperm trees , 2023, IAWA Journal.

[3]  A. Jacobsen,et al.  Vessel diameter polymorphism determines vulnerability-to-embolism curve shape , 2023, IAWA Journal.

[4]  A. Jacobsen,et al.  The functional significance of tracheids co-occurring with vessels in xylem of Eudicots suggests a role in embolism tolerance , 2023, IAWA Journal.

[5]  F. García-Oliva,et al.  Seasonal temperature and precipitation regimes drive variation in the wood of oak species (Quercus) along a climatic gradient in western Mexico , 2023, IAWA Journal.

[6]  S. Jansen,et al.  Functional xylem characteristics associated with drought-induced embolism in angiosperms. , 2022, The New phytologist.

[7]  P. Baas,et al.  Wood Anatomy of Modern and Fossil Fagales in Relation to Phylogenetic Hypotheses, Familial Classification, and Patterns of Character Evolution , 2021, International Journal of Plant Sciences.

[8]  H. Pereira,et al.  Quercus rotundifolia Bark as a Source of Polar Extracts: Structural and Chemical Characterization , 2021, Forests.

[9]  A. Balzano,et al.  Xylem and Phloem Formation Dynamics in Quercus ilex L. at a Dry Site in Southern Italy , 2021, Forests.

[10]  R. Aloni Vascular Differentiation and Plant Hormones , 2021 .

[11]  C. R. Marcati,et al.  Phloem development, growth markers, and sieve-tube longevity in two Neotropical trees , 2020 .

[12]  L. S. Funch,et al.  The growth ring concept: seeking a broader and unambiguous approach covering tropical species , 2019, Biological reviews of the Cambridge Philosophical Society.

[13]  C. R. Marcati,et al.  Duration of cambial activity is determined by water availability while cambial stimulus is day-length dependent in a Neotropical evergreen species , 2017 .

[14]  M. Gloor,et al.  Does Cedrela always form annual rings? Testing ring periodicity across South America using radiocarbon dating , 2017, Trees.

[15]  Susana Valencia-A.,et al.  Diversidad del género Quercus (Fagaceae) en México , 2017 .

[16]  L. Plavcová,et al.  An ecophysiological and developmental perspective on variation in vessel diameter. , 2017, Plant, cell & environment.

[17]  P. Baas,et al.  IAWA List of Microscopic Bark Features , 2016 .

[18]  A. Meyra,et al.  First insights into the functional role of vasicentric tracheids and parenchyma in eucalyptus species with solitary vessels: do they contribute to xylem efficiency or safety? , 2016, Tree physiology.

[19]  C. R. Marcati,et al.  Cambial dormancy lasts 9 months in a tropical evergreen species , 2016, Trees.

[20]  J. Gričar,et al.  Structure and subsequent seasonal changes in the bark of sessile oak (Quercus petraea) , 2015, Trees.

[21]  H. Pereira,et al.  Age trends in the wood anatomy of Quercus faginea , 2014 .

[22]  S. Mayr,et al.  Freeze-Thaw Stress: Effects of Temperature on Hydraulic Conductivity and Ultrasonic Activity in Ten Woody Angiosperms1 , 2013, Plant Physiology.

[23]  I. García‐González,et al.  Comparative cambial dynamics and phenology of Quercus robur L. and Q. pyrenaica Willd. in an Atlantic forest of the northwestern Iberian Peninsula , 2013, Trees.

[24]  H. Pereira,et al.  Bark anatomy and cell size variation in Quercus faginea , 2013, Turkish Journal of Botany.

[25]  S. Carlquist How wood evolves: a new synthesis , 2012 .

[26]  J. Sperry,et al.  Rare pits, large vessels and extreme vulnerability to cavitation in a ring-porous tree species. , 2012, The New phytologist.

[27]  H. Pereira,et al.  Bark anatomy of Quercus cerris L. var. cerris from Turkey , 2011, Turkish Journal of Botany.

[28]  L. Yáñez-Espinosa,et al.  Phenology and radial stem growth periodicity in evergreen subtropical rainforest trees , 2010 .

[29]  H. Pereira,et al.  Characterization of Cork Oak (Quercus Suber) Wood Anatomy , 2009 .

[30]  C. R. Marcati,et al.  SEASONAL VARIATION IN WOOD FORMATION OF CEDRELA FISSILIS (MELIACEAE) , 2006 .

[31]  R. Evert Esau's Plant Anatomy,: Meristems, Cells And Tissues Of The Plant Body- Their Structure, Function And Development , 2005 .

[32]  Sarifuddin,et al.  Tree ring analysis , 1999, Canadian Conference on Electrical and Computer Engineering, 2005..

[33]  R. Tognetti,et al.  Identification, measurement and interpretation of tree rings in woody species from mediterranean climates , 2003, Biological reviews of the Cambridge Philosophical Society.

[34]  O. Dünisch,et al.  FORMATION OF INCREMENT ZONES AND INTRAANNUAL GROWTH DYNAMICS IN THE XYLEM OF SWIETENIA MACROPHYLLA, CARAPA GUIANENSIS, AND CEDRELA ODORATA (MELIACEAE) , 2002 .

[35]  Prof. Dr. Sherwin Carlquist,et al.  Comparative Wood Anatomy , 2001, Springer Series in Wood Science.

[36]  J. Sperry,et al.  The relationship between xylem conduit diameter and cavitation caused by freezing. , 1999, American journal of botany.

[37]  J. Dupouey,et al.  A new attempt at discrimination between Quercus petraea and Quercus robur based on wood anatomy , 1997 .

[38]  S. Davis,et al.  Biophysical Perspectives of Xylem Evolution: is there a Tradeoff of Hydraulic Efficiency for Vulnerability to Dysfunction? , 1994 .

[39]  Dr. Philip R. Larson,et al.  The Vascular Cambium , 1994, Springer Series in Wood Science.

[40]  M. Trockenbrodt Qualitative Structural Changes during Bark Development in Quercus Robur, Ulmus Glabra, Populus Tremula and Betula Pendula , 1991 .

[41]  H. Behnke,et al.  Sieve Elements , 1990, Springer Berlin Heidelberg.

[42]  S. Carlquist Vasicentric Tracheids as a Drought Survival Mechanism in the Woody Flora of Southern California and Similar Regions; Review of Vasicentric Tracheids , 1985 .

[43]  M. Zimmermann Xylem Structure and the Ascent of Sap , 1983, Springer Series in Wood Science.

[44]  A. Ghouse,et al.  Periodicity of Cambium and the Formation of Xylem and Phloem in Mimusops elengi L., an Evergreen Member of Tropical India1)1)Dedicated to the memory of Professor K. A. Chowdhury, the founder of Plant Anatomy Laboratory in Aligarh Muslim University. , 1983 .

[45]  S. Carlquist Ecological strategies of xylem evolution , 1975 .

[46]  D. Aikman Phloem transport , 2020, Nature.

[47]  R. Evert,et al.  Seasonal Cycle of Phloem Development in Woody Vines , 1970, Botanical Gazette.

[48]  R. Evert,et al.  SEASONAL DEVELOPMENT OF THE SECONDARY PHLOEM IN ACER NEGUNDO , 1969 .

[49]  R. Evert,et al.  SEASONAL DEVELOPMENT OF THE SECONDARY PHLOEM IN PINUS , 1968 .

[50]  R. Evert,et al.  Seasonal Development of the Secondary Phloem in Populus tremuloides , 1968, Botanical Gazette.

[51]  R. Evert,et al.  THE CAMBIUM AND SEASONAL DEVELOPMENT OF THE PHLOEM IN ROBINIA PSEUDOACACIA , 1967 .

[52]  H. Fritts Growth-Rings of Trees: Their Correlation with Climate , 1966, Science.

[53]  T. C. Whitmore,et al.  STUDIES IN SYSTEMATIC BARK MORPHOLOGY , 1963 .

[54]  R. Evert THE CAMBIUM AND SEASONAL DEVELOPMENT OF THE PHLOEM IN PYRUS MALUS , 1963 .

[55]  E. M. Gifford,et al.  A staining combination for phloem and contiguous tissues. , 1953, Stain technology.

[56]  E. Artschwager The time factor in the differentiation of secondary xylem and phloem in Pecan. , 1950 .

[57]  K. Esau Phloem structure in the grapevine and its seasonal changes , 1948 .

[58]  S. G. Gilbert,et al.  Evolutionary Significance of Ring Porosity in Woody Angiosperms , 1940, Botanical Gazette.

[59]  W. G. Land Botanical Microtechnique , 1916, Botanical Gazette.